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HARVARD    UNIVERSITY 
-^ 

Library  of  the 

Museum  of 

Comparative  Zoology 


(US  ISSN  0027-4100) 


The  Systematics  of  Neotropical 

Orb-weaving  Spiders  in  the 

Genus  Metepeira  (Araneae:  Araneidae) 


WILLIAM  H.  PIEL 


( 


MCZ 
LIBRARY 

JUL  0  3  2001 


HARVARD 
UNIVERSITY 


HARVARD  UNIVERSITY 

CAMBRIDGE,  MASSACHUSETTS,  U.S.A. 


VOLUME  157,  NUMBER  1 
8  JUNE  2001 


(US  ISSN  0027-4100) 


PUBLICATIONS  ISSUED 

OR  DISTRIBUTED  BY  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 

HARVARD  UNIVERSITY 


Breviora  1952- 

bulletin  1863- 

Memoirs  1865-1938 

JoHNSONiA,  Department  of  Mollusks,  1941-1974 

Occasional  Papers  on  Mollusks,  1945- 

SPEGIAL  PUBLICATIONS. 

1.  Whittington,  H.  B.,  and  W.  D.  I.  Rolfe  (eds.),  1963  Phylogeny  and 
Evolution  of  Crustacea.  192  pp. 

2.  Turner,  R.  D.,  1966.  A  Survey  and  illustrated  Catalogue  of  the  Tere- 
dinidea  (Mollusca:  Bivalvia).  265  pp. 

3.  Sprinkle,  J.,  1973.  Morphology  and  Evolution  of  Blastozoan  Echino- 
derms.  284  pp. 

4.  Eaton,  R.  J.,  1974.  A  Flora  of  Concord  from  Thoreaus  Time  to  the 
Present  Day.  236  pp. 

5.  Rhodin,  A.  G.  J.,  and  K.  Miyata  (eds.),  1983.  Advances  in  Herpetology 
and  Evolutionary  Biology:  Essays  in  Honor  of  Ernest  E.  Williams. 

725  pp. 

6.  Angelo,  R.,  1990.  Concord  Area  Trees  and  Shrubs.  118  pp. 

Other  Publications. 

Bigelow,  H.  B.,  and  W.  C.  Schroeder,  1953.  Fishes  of  the  Gulf  of  Maine. 
Reprinted  1964. 

Brues,  C.T.,  A.  L.  Melander,  and  F.  M.  Carpenter,  1954.  Classification  of 
Insects.  {Bulletin  of  the  M.  C.  Z,  Vol.  108.)  Reprinted  1971. 

Creighton,  W.  S.,  1950.  The  Ants  of  North  America.  Reprinted  1966. 

Lyman,  C.  P.,  and  A.  R.  Dawe  (eds.),  1960.  Proceedings  of  the  First  In- 
ternational Symposium  on  Natural  Mammalian  Hibernation.  {Bulletin 
of  the  M.  C.  Z,  Vol  124.) 

Orinthological  Gazetteers  of  the  Neotropics  (1975-). 

Peter's  Check-list  of  Birds  of  the  World,  vols.  1-16. 

Proceedings  of  the  New  England  Zoological  Club  1899-1947.  (Complete 
sets  only.) 

Proceedings  of  the  Boston  Society  of  Natural  History. 

Price  list  and  catalog  of  MCZ  publications  may  be  obtained  from  Publica- 
tions Office,  Museum  of  Comparative  Zoology,  Harvard  University,  Cambridge, 
Massachusetts  02138,  U.S.A. 

This  publication  has  been  printed  on  acid-free  permanent  paper  stock. 

©  The  President  and  Fellows  of  Harvard  College  2001 . 


THE  SYSTEMATICS  OF  NEOTROPICAL  ORB-WEAVING  SPIDERS  IN 
THE  GENUS  METEPEIRA  (ARANEAE:  ARANEIDAE) 


WILLIAM  H.  PIEU 

CONTENTS 


Abstract  _ ___. 1 

Introduction  2 

Acknowledgments  2 

Materials  and  Methods  3 

Collections  Examined  3 

Locality  Data  Storage  and  Manipulation 4 

Examination  and  Illustration  4 

Metepeira  F.  O.  P.-Cambridge  5 

Key  to  Female  Metepeira  12 

Key  to  Male  Metepeira 17 

Metepeira  foxi  Group  19 

1.  Metepeira  datona  ChamberHn  and  Ivie  20 

2.  Metepeira  desenderi  Baert  21 

3.  Metepeira  grandiosa  grandiosa 
Chamberlin  and  Ivie 23 

4.  Metepeira  grandiosa  alpina 
Chamberlin  and  Ivie  24 

Metepeira  vigilax  Group 26 

5.  Metepeira  cajahamba  New  Species  ....  26 

6.  Metepeira  glornerabilis  (Keyserling)  ...  28 

7.  Metepeira  vigilax  (Keyserling)  30 

8.  Metepeira  rectangula  (Nicolet)  32 

Metepeira  labyrinthea  Group  33 

9.  Metepeira  spinipes  F.  O.  P.-Cambridge  ..  34 

10.  Metepeira  lacandon  New  Species  37 

Metepeira  nigriventris  Group 38 

11.  Metepeira  nigriventris  (Taczanowsld)  38 

12.  Metepeira  tarapaca  New  Species  40 

13.  Metepeira  calamuchita  New  Species  ..  42 

14.  Metepeira  galatheae  (Thorell)  43 

15.  Metepeira  karkii  (Tullgren)  46 

Metepeira  conipsa  Group 47 

16.  Metepeira  compsa  (Chamberlin)  48 

17.  Metepeira  roraima  New  Species  53 

18.  Metepeira  gressa  (Keyserling)  54 

Metepeira  incrassata  Group  56 

19.  Metepeira  maija  New  Species 56 

20.  Metepeira  inca  New  Species  58 


'  Museum  of  Comparative  Zoology,  Harvard  Uni- 
versity, Cambridge,  Massachusetts  02138.  Current 
address:  Institute  of  Evolutionaiy  and  Ecological  Sci- 
ences, Leiden  University,  2311  GP  Leiden,  The 
Netherlands;  piel@rulsfb.leidenuniv.nl. 


21.  Metepeira  gosoga  Chamberlin  and  Ivie 
59 

22.  Metepeira  ohnec  New  Species  60 

23.  Metepeira  comanche  Levi  62 

24.  Metepeira  pimungan  New  Species  62 

25.  Metepeira  triangidaris  (Franganillo)  ..  63 

26.  Metepeira  arizonica  Chamberlin  and 

Ivie  66 

27.  Metepeira  atascadero  New  Species  ....  67 

28.  Metepeira  incrassata  F.  O.  P.- 
Cambridge   68 

Metepeira  ventura  Group  71 

29.  Metepeira  ventura  Chamberlin  and 

Ivie  71 

30.  Metepeira  revillagigedo  New  Species  73 

31.  Metepeira  celestun  New  Species  74 

32.  Metepeira  uncata  F.  O.  P.-Cambridge  ...  76 

33.  Metepeira  crassipes  Chamberlin  and 

Ivie  77 

34.  Metepeira  chilapae  Chamberlin  and 

Ivie  78 

Metepeira  ininima  Group  80 

35.  Metepeira  petatlan  New  Species 80 

36.  Metepeira  minima  Gertsch 82 

37.  Metepeira  pacifica  New  Species 84 

38.  Metepeira  jamaicensis  Archer 86 

Literature  Cited  88 

Index 91 

Abstract.  Of  the  39  species  and  three  subspecies 
of  the  orb-weaver  genus  Metepeira  in  the  Americas, 
36  species  and  two  subspecies  are  known  to  occur 
outside  of  the  U.S.  and  Canada.  Yet,  despite  their 
conspicuous  webs,  diurnal  foraging,  and  relatively 
common  presence,  the  taxonomy  of  Metepeira  is 
poorly  understood,  probably  because  the  genitalia  are 
small  and  difficult  to  distinguish.  In  fact,  many  names 
for  species  south  of  the  U.S.  were,  at  some  time,  in- 
correctly synonymized  with  the  name  Metepeira  la- 
byrinthea. In  this  paper,  14  new  species  are  named 
(Metepeira  atascadero,  M.  cajahamba,  M.  calamuchi- 
ta, M.  celestun,  M.  inca,  M.  lacandon,  M.  maya,  M. 
ohnec,  M.  pacifica,  M.  petatlan,  M.  pimungan,  M.  re- 
villagigedo, M.  roraima,  M.  tarapaca);  11  new  junior 


Bull.  Mus.  Comp.  ZooL,  157(1):  1-92,  June,  2001         1 


2         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


synonyms  are  reported  (M.  acostai,  M.  bani,  M.  dom- 
inicana,  M.  grinnelli,  M.  latigyna,  M.  perezi,  M.  san- 
ta,  M.  salei,  M.  seditiosa,  M.  vaurieorum,  M.  virgi- 
nensis);  five  cases  of  erroneously  synonymized  names 
are  reversed;  22  species  and  two  subspecies  are  re- 
described  (M.  arizonica,  M.  triangidaris,  M.  chilapae, 
M.  Comanche,  M.  compsa,  M.  crassipes,  M.  datona, 
M.  desenderi,  M.  galatheae,  M.  glo7nerabilis,  M.  go- 
soga,  M.  grandiosa  alpina,  M.  grandiosa  grandiosa, 
M.  gressa,  M.  incrassata,  M.  jamaicensis,  M.  karkii, 
M.  minima,  M.  nigriventris,  M.  rectangula,  M.  spi- 
nipes,  M.  uncata,  M.  ventura,  M.  vigilax);  and  a  key 
to  all  Metepeira  species  is  presented.  In  addition,  sev- 
eral ecological  and  life  history  observations  are  re- 
ported for  various  species. 

INTRODUCTION 

The  absence  of  a  comprehensive  revi- 
sion of  Neotropical  Metepeira  has  left  the 
taxonomy  of  this  group  in  shambles.  Over 
the  years,  a  fair  number  of  species  have 
been  named,  particularly  by  A.  F.  Archer, 
R.  V.  Chamberlin,  and  W.  Ivie.  However, 
these  efforts  have  been  sporadic  and,  for 
the  most  part,  scant.  For  example,  the  de- 
scription of  Metepeira  dominicana  (Ar- 
cher, 1965)  provides  little  information  oth- 
er than  "form  typical  of  Metepeira  in  all 
respects,"  a  few  measurements,  and  two 
unrecognizable  figures.  Even  when  species 
are  properly  described  they  have  far  less 
taxonomic  value  when  published  alone,  in 
the  absence  of  a  full  comparative  revision. 

The  poor  understanding  of  Metepeira 
taxonomy  has  persisted  despite  great  eco- 
logical and  behavioral  interest  in  this  ge- 
nus. Indeed,  many  species  are  obligate  or 
facultative  social  species  and  offer  excel- 
lent models  for  investigating  genetic  and 
environmental  factors  that  influence  colo- 
ny formation  (e.g.,  Uetz  and  Cangialosi, 
1986;  Uetz  et  al,  1987).  The  monumental 
work  carried  out  over  many  years  by  G.  W. 
Uetz  has  made  great  strides  in  our  under- 
standing of  gregarious  social  behavior  in 
spiders  and  in  risk-sensitive  foraging  the- 
ory in  general  (e.g.,  Uetz,  1996).  Still,  in 
the  absence  of  solid  taxonomic  literature, 
behavioral  ecologists  have  been  forced  to 
apply  informal  names  to  their  study  ani- 
mals (e.g.,  Metepeira  "atascadero"  in  Uetz 
[1989]  or  Metepeira  "Species  A"  in  Viera 


[1989]),  but  this  practice  can  lead  to  trou- 
ble. In  one  case,  the  behavior  of  several 
different  species  was  initially  studied  un- 
der the  false  assumption  that  they  all  be- 
longed to  the  same  species  (e.g.,  Uetz  et 
al.,  1982).  Clearly,  a  strong  taxonomic 
foundation  is  important  for  further  biolog- 
ical work. 

Ultimately,  the  relatively  small,  indis- 
tinct genitalia  and  the  relatively  homoge- 
neous abdominal  patterns  are  to  blame  for 
the  weakness  in  our  knowledge  of  Mete- 
peira taxonomy.  Many  of  these  species  are 
undoubtedly  hard  to  distinguish,  and  this 
fact  has  surely  intimidated  arachnologists 
from  taking  on  the  painful  task  of  revising 
the  group.  In  the  absence  of  good  distin- 
guishing characteristics,  the  catalogs  of 
Bonnet  (1957)  and  Roewer  (1942)  synon- 
ymized the  names  of  many  Neotropical 
species  with  the  name  Metepeira  lahyrin- 
thea.  Levi's  (1977)  revision  of  Nearctic 
species  observes  that  M.  labyrinthea  is  ac- 
tually limited  to  the  eastern  United  States. 
One  task  in  this  revision  consists  of  reas- 
serting the  names  of  species  that  were  im- 
properly synonymized  and  clarifying  the 
diagnostic  characters  that  are  needed  to 
identify  them. 

ACKNOWLEDGMENTS  i 

This  paper  is  part  of  my  Ph.D.  thesis  for 
the  Department  of  Organismic  and  Evo- 
lutionary Biology,  Harvard  University.  I  ain 
indebted  to  many  people  for  their  help, 
assistance,  and  encouragement  in  this  pro- 
ject. I  am  especially  thankful  for  the  ded- 
ication and  support  of  my  advisors,  Her- 
bert W.  Levi  and  Edward  O.  Wilson.  I  am 
grateful  that  my  colleagues  in  the  Depart- 
ment of  Invertebrate  Zoology  provided 
such  a  pleasant  place  to  work:  Edward 
Cutler,  Ardis  Johnston,  Laura  Leibensper- 
ger,  Damhnait  McHugh,  Diana  Sherry, 
Van  Wallach,  and  Dee  Woessner,  among 
others. 

Field  collecting  and  new  specimen  ac- 
quisitions were  made  possible  with  the 
help  of  Gita  Bodner,  Fundacion  Capacitar, 
Tim  Coonan  (CINP),  Fred  Coyle,  Dawn 


Metepeira  •  Piel 


Fitzpatrick,  Germania  Jacome,  Antdnia 
Monteiro,  Tila  Perez,  George  Putnam, 
Linda  Rayor,  Grace  Smith  (NAWF),  and 
George  Uetz.  I  am  particularly  indebted  to 
George  Uetz  for  his  assistance  and  corre- 
spondence. 

I  am  thankful  for  the  comments  by 
those  who  read  this  paper — especially  to 
the  members  on  my  thesis  committee:  H. 
W.  Levi,  N.  E.  Pierce,  and  E.  O.  Wilson. 
I  am  also  indebted  to  Kathy  Horton  for 
her  help  in  formatting  and  preparing  the 
manuscript  and  to  the  Colles  Fund  for  de- 
fraying the  costs  of  publication.  Curators 
at  various  institutions  who  lent  me  speci- 
mens are  listed  in  the  Materials  and  Meth- 
ods section.  I  cannot  overstress  the  value 
of  museum  collections  and  expert  curators, 
without  which  research  in  taxonomy  would 
not  be  possible.  Museum  collections  are 
the  most  important  tools  available  for  un- 
derstanding biodiversity. 

MATERIALS  AND  METHODS 

Collections  Examined.  The  taxonomic 
revision  was  carried  out  on  specimens  bor- 
rowed from  the  following  collections.  The 
abbreviations  correspond  to  those  listed 
with  each  record  after  eveiy  species  de- 
scription. I  am  grateful  to  the  museums, 
curators,  and  staff  that  graciously  loaned 
the  material. 

ADC  A.  Dean,  Texas  A&M  University, 

College  Station,  Texas,  United 
States 

AMNH  American  Museum  of  Natural 
History,  New  York,  United 
States;  N.  Platnick,  L.  Sorkin 

BMNH  Natural  History  Museum,  Lon- 
don, England;  P.  Hillyard 

CAS  California  Academy  of  Sciences, 

San  Francisco,  California,  Unit- 
ed States;  C.  Griswold 

CV  Carlos   Valderrama  A.;    Bogota, 

Colombia 

FSCA  Florida  State  Collection  of  Ar- 
thropods, Gainesville,  Florida, 
United  States;  G.  B.  Edwards 

IRSNB      Institut  Royal  des  Sciences  Na- 


turelles  de  Belgique,  Brussels, 
Belgium;  L.  Baert 

JAK  J.  A.  Kochalka,  Ciudad  Univer- 

sitaria,  Paraguay 

JEC  J.    Carico,   Lynchburg,  Virginia, 

United  States 

JMM          J.  Maes,  Leon,  Nicaragua 

MACN  Museo  Argentino  de  Ciencias 
Naturales,  Buenos  Aires,  Argen- 
tina; E.  A.  Maury,  C.  L.  Scioscia 

MCN  Museu  de  Ciencias  Naturais, 
Fundagao  Zoobotanica  do  Rio 
Grande  do  Sul,  Porto  Alegre, 
Rio  Grande  do  Sul,  Brazil;  E.  H. 
Buckup,  M.  A.  L.  Marques 

MCZ  Museum  of  Coniparative  Zool- 

ogy, Harvard  University,  Cam- 
bridge, Massachusetts,  United 
States;  H.  W.  Levi 

MECN  Museo  Ecuatoriano  de  Ciencias 
Naturales,  Quito,  Ecuador;  Ger- 
mania Estevez  Jacome 

MEG  M.  E.  Galiano,  Buenos  Aires, 
Argentina 

MLJC  Maria  Luisa  Jimenez,  Centro  de 
Investigaciones  Bioldgicas  del 
Noroeste,  La  Paz,  Mexico 

MLP  Museo  de  Universidad  Nacional, 

La  Plata,  Argentina;  R.  F.  Arro- 
zpide,  C.  Sutton 

MNRJ  Museu  Nacional,  Rio  de  Janeiro, 
Brazil;  A.  Timotheo  da  Costa 

MNSD  Museo  Nacional  de  Historia 
Natural,  Santo  Domingo,  Re- 
publica  Dominicana;  Felix  Del 
Monte 

MUSM  Museo  de  Historia  Natural, 
Universidad  Nacional  Mayor  de 
San  Marcos,  Lima,  Peru;  D.  Silva 

MZSP  Museu  de  Zoologia,  Universida- 
de  de  Sao  Paulo,  Sao  Paulo,  SP, 
Brazil;  P.  Vanzolini,  J.  L.  Leme 

MZUF  Museo  Zoologico  de  "La  Spe- 
cola"  Universita  di  Firenze, 
Florence,  Italy;  S.  Whitman 

NRMS  Naturhistoriska  Riksmuseet, 
Stockholm,  Sweden;  T.  Krones- 
tedt 

PAN  Polska  Akademia  Nauk,  Warsza- 


4         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


wa,  Poland;  J.  Proszynski,  A.  Slo- 
jewska,  W.  B.  Jedryczkowski 

REL  R.  E.  Leech,  Edmonton,  Alber- 

ta, Canada 

SMF  Forschungsinstitut      Sencken- 

berg,  Frankfurt  am  Main,  Ger- 
many; M.  Grasshoff 

SR  Susan  Riechert,  Knoxville,  Ten- 

nessee, United  States 

USNM  National  Museum  of  Natural 
History,  Smithsonian  Institution, 
Washington,  D.C.,  United 
States;  J.  Coddington,  S.  F. 
Larcher 

ZMB  Zoologisches       Museum      der 

Humboldt  Universitat,  Berlin, 
Germany;  M.  Moritz 

ZMUC  Zoologisk  Museum,  Gopenha- 
gen,  Denmark;  H.  Enghoff,  N. 
Scharff 

ZSM  Zoologische    Staatssammlung, 

Munich,  Germany 

Locality  Data  Storage  and  Manipula- 
tion. Locality  data  from  each  collection  vial 
were  entered  into  a  database  designed  us- 
ing Glaris  FileMaker  Pro®.  Geographic 
coordinates  were  added  to  locality  data 
that  lacked  them  using  maps,  USBGN  gaz- 
etteers, and  on-line  databases  (http://164. 
214.2.59/gns/html/  and  http://mapping. 
usgs.gov/www/gnis/).  Occasionally  locality 
information  was  illegible  or  unknown  or 
one  of  several  homonymous  sites.  In  such 
cases  a  reasonable,  educated  guess  was 
inade  and  a  "[?]"  designation  was  append- 
ed to  the  locality.  In  some  cases  the  itin- 
erary of  a  collector  was  reconstructed  from 
other  known  records,  and  the  ambiguous 
locality  was  assigned  a  coordinate  halfway 
between  the  previous  and  following  known 
collection  sites.  The  locality  database 
worked  in  concert  with  the  mapping  pro- 
gram Atlas  Pro®  to  generate  thematic 
maps  on  the  fly.  These  maps  helped  in  the 
process  of  delimiting  species  and  discov- 
ering cryptic  species. 

Elevation  (in  meters)  was  estimated  for 
each  locality  that  lacked  this  information. 
In  some  cases,  elevation  was  estimated  us- 


ing contour  maps,  such  as  DMAAG  ONG 
aeronautical  maps;  in  most  cases,  elevation 
was  estimated  using  NOAA  data  with  an 
on-line  database  server  (http://phylogeny. 
harvard.edu/~piel/find.htiril). 

The  enhanced  locality  database  was 
used  to  reveal  ecological  and  life  history 
traits.  Seasonality  of  species  was  expressed 
by  plotting  a  circular  histogram  showing 
the  relative  amount  of  collecting  activity 
per  5-day  interval  (Figs.  300-337).  While 
locality  dates  alone  cannot  control  for  the 
seasonal  activity  of  human  collectors,  these 
data  at  least  provide  an  estimate  of  spider 
seasonal  abundance,  if  only  approximate. 
Some  syiupatric  species  show  incongruous 
seasonal  abundance,  which  is  at  least  some 
evidence  that  seasonality  of  spider  collec- 
tors does  not  unduly  overshadow  the  sea- 
sonality of  the  spiders  themselves. 

Examination  and  Illustration.  Speci- 
mens were  examined  under  80%  ethanol 
in  a  dish  with  light  and  dark  sand  grains 
for  specimen  support.  Digital  photographs 
of  preserved  specimens  were  taken 
through  a  Nikon  SMZ-10  photomicro- 
scope  using  a  Panasonic  WV-CL320  GGD 
video  camera,  chosen  for  its  high  sensitiv- 
ity to  light.  Video  images  were  captured 
using  a  Quicklmage®24  digitizer  and  ed- 
ited on  a  Quadra  700  Macintosh®  com- 
puter. The  computer  allows  relatively  in- 
expensive pictures  to  be  printed  rapidly  on 
a  1,200  dpi  Xante®  Accel-a- Writer  8200 
laser  printer.  Digital  pictures  were  used  to 
help  sort  out  individuals  to  species,  to  cre- 
ate publishable  pictures  of  gross  dorsal 
and  ventral  markings,  and  to  aid  in  the  il- 
lustration of  genitalia.  As  an  aid  in  illustra- 
tion, the  digital  pictures  functioned  as  a 
camera  lucida  because  they  assured  accu- 
racy when  drawing  the  proportions  of  gen- 
ital parts  and  scle rites.  Usually  a  digital 
picture  was  laid  over  carbon  paper  and  an 
outline  of  the  genitalia  was  transferred  to 
coquille  board  underneath.  The  illustra- 
tion continued  on  the  coquille  board  using 
a  Staedtler  OmniGhrom®  pencil  and  a 
drafting  pen  with  India  ink  and  then  was 
scanned  at  600  dpi  on  a  LaGie  Silverscan- 


Metepeira  •  Piel 


ner  II®.  The  resulting  digital  image  was 
edited  in  Adobe  Photoshop®  and  reduced 
in  size  to  1,200  dpi.  The  edited  figures 
were  finally  arranged  on  plates  using  Can- 
vas®. 

External  genital  structures  were  manip- 
ulated with  pins  to  reveal  hidden  parts. 
The  terminal  division  on  the  male  palp  is 
hinged,  so  it  had  to  be  pried  open  to  see 
the  embolus  and  embolic  apophyses  prop- 
erly. In  females,  mating  plugs  had  to  be 
removed  from  epigynal  openings  using 
pins.  Sometimes  the  entire  epigynum  was 
partly  cut  from  the  body  so  as  to  see  it 
from  a  posterior  view. 

Internal  genital  structures  were  studied 
by  clearing  them  in  clove  oil  and  examin- 
ing them  using  an  Olympus  BH-2  com- 
pound microscope.  Sketches  were  made 
directly  on  the  computer  in  Canvas®  by 
aiming  the  camera  lucida  at  the  computer 
monitor.  While  internal  genital  structures 
helped  in  the  process  of  delimiting  spe- 
cies, they  did  not  prove  to  be  as  useful  as 
external  genital  structures  in  describing 
species;  thus,  these  working  sketches  are 
not  figured  herein. 

Measurements  of  the  spiders  were  tak- 
en using  a  Leitz  stereo  dissecting  micro- 
scope with  a  calibrated  reticule.  Sizes  of 
leg  articles,  eyes,  and  carapace,  were  per- 
formed on  one  specimen  of  each  sex,  for 
each  species.  The  respective  localities  of 
the  candidate  specimens  were  indicated  in 
the  descriptions.  This  study  placed  little 
reliance  on  spider  leg  measurements  be- 
cause they  are  not  usually  very  useful  in 
spider  taxonomy,  and  because  Metepeira 
species  are  notorious  for  their  variability  in 
size  (Levi,  1977;  Piel,  1996). 

All  eye  sizes  were  reported  as  a  ratio  of 
the  posterior  median  eye  diameters  to  the 
diameter  of  every  other  eye  type.  For  ex- 
ample, in  the  case  of  "ratio  of  eye  diame- 
ters: posterior  medians  and  anterior  me- 
dians 2.0,  anterior  laterals  0.5,  posterior 
laterals  1.0,"  the  reader  should  interpret 
the  anterior  medians  to  be  half  the  size  of 
the  posterior  medians,  and  the  anterior  lat- 
erals to  be  twice  the  size  of  the  posterior 


medians.  Eye  separations  were  expressed 
in  terms  of  their  own  diameters,  or  in 
terms  of  the  anterior  lateral  eyes  when  be- 
tween eyes  of  different  types.  Oval  eyes 
were  measured  as  an  average  of  the  lon- 
gest and  shortest  lengths. 

In  parallel  with  the  last  revision  of  Me- 
tepeira (Levi,  1977),  leg  measurements 
were  made  on  each  article  distal  to  the  tro- 
chanter for  the  first  leg  and  on  the  com- 
bined lengths  of  the  patellae  and  tibiae  for 
all  remaining  legs.  Variation  in  total  body 
size  was  provided  as  an  average,  minimum, 
and  maximum  of  the  total  lengths  from  a 
number  of  mature  specimens,  usually  cho- 
sen from  a  wide  geographic  spread. 

Metepeira  F.  O.  P. -Cambridge 

Metepeira  F.  O.  P.-Cambridge,  1903:  457.  Type  spe- 
cies by  original  designation  M.  spinipes  F.  O.  P.- 
Cambridge 1903.  The  name  is  feminine. 

Diagnostic  Abstract.  Web  combines  bar- 
rier or  scaffolding  structure  surrounding  a 
classic  araneid  orb  with  a  retreat  suspend- 
ed in  air  (Fig.  1).  Like  a  raccoon  with  its 
facial  colors  reversed,  the  eye  region  is 
lighter  than  any  other  part  of  the  carapace 
(Fig.  2).  The  venter  has  a  wide  median 
white  line  set  on  a  black  background  that, 
with  only  some  exception,  extends  anteri- 
orly on  the  sternum  (Fig.  3).  With  one  ex- 
ception, the  total  lengths  of  distal  leg  ar- 
ticles (metatarsus  and  tarsus)  exceed  that 
of  the  middle  articles  (patella  and  tibia). 
The  median  apophysis  has  two  distinctive 
flagella  (F  in  Fig.  5)  and,  in  some  species, 
an  easily  recognizable  keel  (K  in  Fig.  5). 
The  dorsal  abdominal  markings  (the  foli- 
um) look  like  an  inverted  fleur-de-lis,  al- 
lowing easy  recognition  of  the  genus  in  the 
field  (Fig.  2). 

Description  and  Diagnosis.  For  field 
ecologists,  the  most  obvious  and  distinctive 
feature  of  Metepeira  is  the  combination  of 
orb  and  barrier  web  (Fig.  1).  The  barrier 
\^eh  forms  scaffolding  around  an  almost 
vertical  orb  and  supports  the  spiders  re- 
treat, which  is  thus  suspended  away  from 
any  substrate. 

In  contrast  to  most  araneids,  the  cara- 


6         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


pace  of  Metepeira  is  lightest  in  the  eye  re- 
gion. However,  this  distinctive  feature 
varies  within  the  genus:  in  the  case  of  M. 
rectangula  (Nicolet,  1849),  the  hghter  re- 
gion takes  up  ahiiost  half  the  carapace 
(Fig.  65);  in  the  case  of  Metepeira  F.  O. 
P.-Cambridge,  1903,  the  lighter  region  is 
usually  limited  to  the  anterior  edges  of  the 
carapace  (Fig.  2).  White,  downy  hairs  of- 
ten cover  the  carapace  but  are  especially 
white  and  conspicuous  on  the  lighter  parts 
of  the  carapace  outside  the  eye  region.  In 
some  species,  such  as  M.  spinipes,  these 
hairs  make  the  carapace  look  gray  or  sil- 
very when  the  spider  is  alive,  but  dark 
brown  when  the  spider  is  in  ethanol. 

The  eyes  of  Metepeira  are  not  particu- 
larly unusual.  Eye  separations  relative  to 
eye  diameters  increase  with  spider  size: 
larger  spiders  tend  to  have  relatively  great- 
er eye  separations.  In  either  sex,  the  pos- 
terior median  eyes  are  between  1.1  and 
1.7  times  the  size  of  the  anterior  medians, 
and  the  separation  between  posterior  me- 
dian eyes  is  between  0.4  and  0.7  of  that 
between  anterior  median  eyes.  The  sepa- 
ration between  the  anterior  median  eyes 
and  the  anterior  lateral  eyes  is  between  1 
and  3.7  times  the  size  of  anterior  median 
eyes  in  males  and  between  one  and  five 
times  the  size  in  females.  The  diameter  of 
the  anterior  median  eyes  exceeds  the 
height  of  the  clypeus. 

The  shape  of  the  female  abdomen  rang- 
es from  wider  than  long  and  rhomboid  (M. 
datona.  Fig.  12)  to  roundish  (e.g.,  M.  de- 
senderi.  Fig.  20;  M.  rectangula.  Fig.  65), 
to  longer  than  wide  and  oval  (e.g.,  M.  inca. 
Fig.  169).  The  dorsal  folium  has  a  recog- 
nizable white  fleur-de-lis  pattern,  usually 
on  a  dark  background,  its  edges  shaped  by 
a  wavy,  zig-zag  white  outline  (Fig.  2).  The 
dorsum  of  live  spiders  is  often  more  red- 
dish— a  pigment  that  rapidly  dissolves  in 
alcohol. 

Somewhat  less  common  among  other 
araneids  is  the  median  white  line  on  the 
venter  of  the  abdomen  (Fig.  3),  which  is 
present  (though  shortened)  even  in  the 
most  darkly  pignriented  species.  However, 


unique  among  araneid  genera  is  the  com- 
bination of  median  white  line  on  the  ven- 
ter and  median  white  line  on  a  black  or 
brown  sternuin.  Some  Metepeira  species 
lack  a  complete  white  line  on  the  sternum, 
but  even  those,  such  as  M.  datona,  that 
usually  have  an  entirely  black  sternum 
nonetheless  sho\v  hints  of  white  markings 
in  some  specimens.  Characteristics  found 
in  the  carapace,  abdomen,  and  sternum  of 
Metepeira  are  also  found  in  Araneus 
koepckeorurn  Levi,  but  this  last  species 
lacks  the  white  line  on  the  venter. 

With  the  exception  of  M.  datona,  and  in 
some  cases,  M.  desenderi,  all  Metepeira 
species  have  a  combined  metatarsus  and 
tarsus  that  is  longer  than  the  combined  pa- 
tella and  tibia.  This  feature  is  unusual 
among  araneids  and  is  not  found  in  Kaira 
O.  P. -Cambridge  or  other  likely  relatives 
to  Metepeira  (Levi,  1977;  Piel  and  Nutt, 
1997). 

In  most  species  the  leg  articles  are 
ringed,  usually  with  brownish  black  on  the 
distal  and  dorsal  surfaces  of  each  article, 
except  for  the  patellae  and  tarsi  which  are 
usually  entirely  dark.  In  mainly  tropical 
and  high-altitude  species,  the  coxae  are 
mostly  black  (e.g..  Fig.  75),  but  in  desert/ 
mesquite  species  they  appear  yellowish 
white  (e.g..  Fig.  28).  I 

Unlike  many  other  araneids — and  per- 
haps because  of  the  small  male  size — the 
coxa  on  leg  I  of  male  Metepeira  lacks  the 
hook  and  corresponding  groove  typically 
found  on  femur  II.  In  addition,  males  lack 
a  tooth  on  the  lateral  side  of  the  endite, 
and  they  lack  a  basal  tooth  on  the  palpal 
femur.  The  phylogenetic  analysis  of 
Scharff  and  Coddington  (1997)  incorrectly 
codes  Metepeira  as  having  a  tooth  on  the 
endite.  However,  had  the  authors  coded 
this  character  as  absent,  they  would  have 
decreased  the  length  of  their  preferred 
tree  because  the  nearest  relatives  hypoth- 
esized for  Metepeira  {Kaira,  Zijgiella,  and 
Singa)  also  lack  this  tooth. 

Macrosetae  usually  concentrate  on  arti- 
cles that  contact  other  spiders  during  mat- 
ing or  grappling.  In  contrast  to  most  gen- 


Metepeira  •  Piel        7 


Barrier  Web 


White  Dorsal  Light  Eye  Region 

Fleur-de-Lis  /  ^^  Median  White 

Pattern        x AW*  ^Ma    Line  on  Sternum 


Orb  Web 


Median  White 
Line  on  Black 
Venter 


vigilax  nigriventris  incrassata  minima 

foxi  labyrinthea  compsa  vcntura 


Figure  1.  Web  of  immature  Metepeira  grandiosa  alpina  from  Chihuahua,  Mexico. 

Figure  2.  Dorsum  of  adult  female  Metepeira  crassipes. 

Figure  3.  Venter  of  adult  female  Metepeira  tarapaca  new  species. 

Figure  4.  Hypothetical  phylogenetic  relationships  among  Metepeira  species  groups.  Shaded  branches  indicate  species  groups 

that  live  in  South  America;  open  branches  indicate  species  groups  that  live  in  North  America,  Central  America,  and  the  Caribbean. 

Abbreviations:  DEA,  distal  embolic  apophysis;  K,  keel  of  median  apophysis;  TA,  terminal  apophysis;  (+),  character  state  gain; 

(-),  character  state  loss. 

Figures  5,  6.  Male  palpus.  5,  mesal  view,  Metepeira  compsa.  6,  ventral  view  of  distal  embolic  division,  Metepeira  labyrintliea 

(Hentz). 

Abbreviations:  BEA,  basal  embolic  apophysis;  C,  conductor;  DEA,  distal  embolic  apophysis;  E,  embolus;  F,  flagellum  on  median 

apophysis;  K,  keel  of  median  apophysis;  MA,  median  apophysis;  TA,  terminal  apophysis;  TO,  terminal  division. 

Figures  7-13.  Metepeira  datona  Chamberlin  and  Ivie  (sp.  1;  17°53'N,  76°19'W).  7,  male  palpus,  mesal.  8,  epigynum,  posterior. 

9,  epigynum,  ventral.  10,  male,  dorsal.  11,  male,  ventral.  12,  female,  dorsal.  13,  female,  ventral. 

Scale  bar:  dorsum  and  venter  figures  1 .0  mm. 


8         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


era  related  to  Araneus,  Metepeira  has  con- 
centrated macrosetae  on  femur  I  instead 
of  tibia  II  (Scharff  and  Coddington,  1997). 
Female  Metepeira  have  between  two  and 
five  macrosetae  on  die  anterior  side  of  the 
femur,  and  between  zero  and  seven  on  the 
anteroventral  side.  Males  typically  have 
inore  setae  than  their  conspecific  females: 
four  to  nine  on  the  anterior  side  and  two 
to  nine  on  the  anteroventral  side.  Variation 
in  the  number  of  macrosetae  appears  to 
correlate  with  body  size.  In  most  species, 
the  inale  palpal  tibia  and  patella  each  have 
two  strong  macrosetae  (Levi,  1977,  fig.  8). 

Compared  with  other  araneid  genera, 
Metepeira  have  rather  small  and  similar 
genitalia,  which  on  the  one  hand  makes 
the  genus  easy  to  recognize,  but  on  the 
other  hand  makes  species  tough  to  iden- 
tify. The  small  epigynum  is  fleshy,  variable 
in  shape,  and  weakly  sclerotized.  Unlike 
Araneus,  Metepeiras  scape  never  has  a 
pocket  but  always  ends  with  a  pointed  tip 
(e.g..  Fig.  31).  The  cleared  epigynum — 
and  in  many  cases  the  uncleared  epigyn- 
um— reveals  a  pair  of  sclerotized  spherical 
structures  where  the  embolus  is  inserted, 
as  well  as  ducts  to  pass  semen  to  the  larger, 
spherical  seminal  receptacles.  In  some 
species,  these  spherical  structures  are  wide 
apart  (e.g..  Figs.  16,  17),  in  others  they  are 
tubular  (e.g..  Figs.  39,  40),  but  in  many, 
they  are  closer  together  (e.g..  Figs.  93,  94). 
Frequently  the  deeper,  large  seminal  re- 
ceptacles can  be  seen  through  uncleared 
tissue  (e.g..  Figs.  201,  295). 

The  male  palp  is  more  distinctive.  In 
particular,  the  median  apophysis  (MA  in 
Fig.  5),  while  not  always  a  good  character 
for  separating  closely  related  species,  is  ex- 
cellent when  it  comes  to  identifying  the 
genus.  Two  flagellae  (F  in  Fig.  5)  grace- 
fully curve  off  the  base  of  the  median 
apophysis,  and  in  some  species,  a  toothed 
or  smooth  keel  (K  in  Fig.  5)  extends  in  the 
opposite  direction.  This  design  is  also  seen 
in  Kaira,  Aculepeira,  and  Amazonepeira, 
but  none  of  these  have  flagellae  that  ap- 
pear so  integral  to  the  base  structure. 

The  tenninal  division  on  the  Metepeira 


palp  is  very  similar  in  almost  all  species. 
When  this  structure  is  pulled  up,  a  basal 
embolic  apophysis  (also  known  as  an  em- 
bolar  lamella)  can  be  seen  in  the  shape  of 
a  club  or  spatula  (E  in  Figs.  5,  6).  Soiue- 
times  a  distal  embolic  apophysis  can  be 
seen  if  it  is  not  hidden  from  view  by  an 
overhanging  terminal  apophysis.  When  the 
terminal  apophysis  is  large  and  sclero- 
tized— which  is  die  case  in  all  but  the  Me- 
tepeira foxi  species  group — it  has  a  rec- 
ognizable toothed  notch,  like  the  mouth 
on  a  wrench  (Fig.  6).  Virgin  males  have  a 
cap  on  the  embolus  that  remains  in  the 
epigynum  after  mating  and  presumably 
serves  as  a  barrier  to  subsequent  mating 
(Levi,  1977).  The  shape  of  the  embolus 
cap  varies  from  tiny  (e.g..  Fig.  178)  to 
short  but  wide  (e.g.,  Fig.  199)  to  large  and 
winged  (Fig.  46).  Finally,  the  terminal  di- 
vision lacks  a  stipes — a  sclerite  between 
the  radix  and  the  embolus  that  is  frequent- 
ly found  in  other  genera  related  to  Ara- 
neus (Scharff  and  Coddington,  1997). 

Natural  History.  All  Metepeira  species 
build  a  unique  web  that  combines  an  orb 
with  a  barrier  web  (Levi,  1977;  Lopez, 
1993).  As  with  Cijrtophora  Simon  or  Me- 
cynogea  Simon  (Levi,  1997),  the  retreat  of 
Metepeira  hangs  in  the  air,  away  from  sub- 
strate, and  is  suspended  by  a  scaffolding 
structure  created  by  the  barrier  web  (Fig. 
1).  The  spider  detects  vibrations  in  the 
web  and  gains  quick  access  to  the  hub  us- 
ing a  signal  line  that  runs  from  the  retreat 
to  the  center  of  the  orb  (Fig.  1).  Tan  col- 
ored egg  sacs  are  strung  together,  usually 
above  the  retreat,  and  the  most  recently 
laid  eggs  are  nearest  to  the  spider.  In  some 
species  the  egg  sacs  and  retreat  are  deco- 
rated with  insect  parts  (e.g.,  M.  spinipes); 
in  other  species  they  are  carefully  wrapped 
by  leaves  and  woven  together  (e.g.,  M.  da- 
tona).  Unlike  the  webs  oi  Cijrtophora  and 
Mecynogea,  the  orb  web  of  Metepeira  is 
oriented  vertically,  and  the  number  of  radii 
and  sticky  spirals  are  more  typical  of  other 
araneines. 

In  some  species,  such  as  M.  pimungan 
(personal  observation)  and,  to  a  lesser  de- 


Metepeira  •  Piel 


gree,  M.  incrassata  (G.  Uetz,  personal 
communication),  juveniles  and  adults 
without  eggs  will  live  on  webs  lacking  a 
suspended  retreat.  Instead,  the  spider  sits 
on  a  white  disk-shaped  stabilimentum  in 
the  center  of  the  hub.  Of  110  M.  pimun- 
gan  specimens  observed  on  San  Miguel  Is- 
land, about  40%  occupied  webs  of  this 
type.  In  two  cases  the  disk  stabilimenta 
were  partly  separated  from  the  hub  by 
barrier  web  lines  and  were  further  bent 
over  to  form  a  partly  covered  protective 
retreat  for  the  spider.  This  observation 
makes  it  possible  to  imagine  that  the  disk 
stabilimentum  seen  in  M.  pimiingan  re- 
sults from  the  fusion  of  the  suspended  re- 
treat with  the  hub. 

When  food  supplies  are  plentiful,  spi- 
ders of  all  kinds  show  an  increased  toler- 
ance for  one  another  and  an  increased  ten- 
dency to  aggregate  (e.g.,  Gillespie,  1987; 
Rypstra,  1986).  The  suspended  retreats 
and  barrier  webs  of  Metepeira,  Cyrtopho- 
ra,  and  Mecynogea  may  actually  further  fa- 
cilitate in  the  formation  of  aggregations  by 
easing  dependency  on  substrate  availabili- 
ty and  by  providing  a  common  support  sys- 
tem (Burgess  and  Witt,  1976;  Uetz,  1986). 
In  any  case,  colony  formation  is  known  to 
occur  in  all  three  genera  (e.g.,  Rypstra, 
1979),  but  especially  in  Metepeira.  Small 
colonial  aggregations  of  two  to  10  individ- 
uals occur  in  M.  datona  (Spiller  and 
Schoener,  1989),  M.  minima  (personal  ob- 
servation), M.  glomerabilis  (R.  Baptista, 
personal  communication),  and  M.  atascad- 
ero  new  species  (e.g.,  Uetz  and  Hodge, 
1990).  Medium-size  colonies  of  10  to  30 
individuals  occur  in  M.  pimungan  (person- 
al observation),  M.  gressa  (Viera  and  Cos- 
ta, 1988),  M.  nigriventris  (L.  Rayor,  per- 
sonal coinmunication),  M.  tarapaca  (V. 
Roth,  locality  label),  and  M.  spinipes  (e.g., 
Uetz,  1988a).  Large  colonies,  sometimes 
in  the  thousands  of  individuals,  commonly 
occur  in  M.  incrassata  (e.g.,  Uetz  and 
Hodge,  1990).  Near  rivers  and  in  other 
lush  habitats,  M.  tarapaca  colonies  can 
reach  200  individuals  (M.  Roy,  personal 
communication).  These  cases  of  social  be- 


havior, broadly  spread  across  seven  differ- 
ent species  groups,  may  mean  that  aggre- 
gation is  a  frequently  lost  and  relatively  old 
trait,  or  it  may  mean  that  species  are  prone 
to  converge  and  evolve  the  same  behavior 
independently. 

Either  way,  much  research  has  focused 
on  elucidating  the  selective  forces  behind 
colonial  behavior  in  Metepeira.  In  partic- 
ular, Uetz  (1988a,b,  1996)  has  provided 
strong  support  for  the  hypothesis  that  Me- 
tepeira forage  using  a  risk-sensitive  strat- 
egy. He  suggests  that  spiders  in  abundant 
habitats  seek  to  minimize  individual  vari- 
ance in  prey  capture  by  aggregating  in  col- 
onies, whereas  spiders  in  poor  habitats 
seek  to  maximize  variance  by  living  soli- 
tarily— perhaps  in  a  risky  attempt  to  find 
areas  of  local  prey  maxima.  The  diversity 
of  social  tendencies  among  species  is 
therefore  commensurate  of  the  diversity  of 
ecological  habitats  that  they  inhabit. 

Indeed,  Metepeira  species  thrive  in  a 
wide  array  of  habitats,  though  often  they 
are  quite  harsh.  These  include  wet,  mon- 
tane cloud  forests  in  Mexico  and  Panama 
(M.  incrassata,  M.  olmec);  tropical  and  wet 
agricultural  areas  (M.  uncata,  M.  vigilax, 
M.  glomerabilis,  M.  roraima);  high-eleva- 
tion pine  forests  (M.  lacandon,  M.  nigri- 
ventris, M.  grandiosa  alpina);  Canadian 
bogs  (M.  grandiosa  palustris);  deciduous 
forests  in  the  eastern  U.S.  (M.  labyrin- 
thea);  Caribbean  coastal  shrubbery  (M.  da- 
tona, M.  minima,  M.  triangularis,  M.  ja- 
maicensis,  M.  maya,  M.  celestun);  Mexican 
mesquite  grasslands  (M.  atascadero,  M. 
chilapae);  Patagonian  dunes  and  scrub  (M. 
galatheae)  and  pampas  grass  (M.  karkii); 
diy  Californian  buckwheat  and  sage  (M. 
crassipes,  M.  ventura,  M.foxi,  M.  grandio- 
sa grandiosa);  and  arid  and  semiarid  de- 
serts (M.  arizonica,  M.  inca,  M.  ventura, 
M.  crassipes).  Although  some  species  (e.g., 
M.  galatheae,  M.  spinipes,  M.  cornpsa)  cov- 
er vast  geographic  areas  and  live  in  many 
different  habitats,  many  species  are  more 
biogeographically  restricted.  In  fact,  sev- 
eral species  follow  narrow  ecological  zones 
that  decrease   in  elevation  with  distance 


10         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No. 


from  the  equator  (e.g.,  M.  rectangula,  M. 
vigilax,  M.  cajahamha.  Fig.  36;  M.  arizon- 
ica.  Fig.  213). 

Close  cohabitation  with  different  inter- 
and  intrageneric  species  is  not  uncommon. 
Colonies  of  M.  incrassata  are  known  to 
contact  webs  of  Nephila  clavipes  Linnaeus 
(Hodge  and  Uetz,  1996)  and  Mecijnogea 
ocosingo  and  Gasteracantha  cancriformis 
(personal  observation).  Often  M.  crassipes, 
M.  Ventura,  M.  foxi,  and  M.  grandiosa 
grandiosa  are  collected  together  (Levi, 
1977),  as  are  M.  minima  and  M.  celesiun 
(personal  observation).  Species  that  have 
been  collected  from  identical  localities, 
though  not  necessarily  at  the  same  time, 
include:  M.  chilapae  and  M.  spinipes;  M. 
chilapae  and  M.  atascadero;  M.  karkii  and 
M.  galatheae;  M.  calamuchita  new  species, 
M.  gressa,  and  M.  galatheae;  M.  rectan- 
gula, M.  calamuchita,  and  M.  galatheae; 
M.  compsa  and  M.  gressa;  M.  vigilax  and 
M.  compsa;  M.  glomerahilis  and  M.  vigilax; 
M.  compsa  and  M.  glotnerabilis;  M.  comp- 
sa and  M.  nigriventris;  M.  compsa  and  M. 
mcfl;  M.  datona  and  M.  jamaicensis;  and 
M.  datona  and  M.  triangularis. 

Despite  the  wide  biogeographic  ranges 
of  M.  compsa  (Puerto  Rico  and  south  to 
Argentina,  Map  8)  and  M.  datona  (His- 
paniola  and  north  to  Florida,  Map  1),  they 
nonetheless  come  geographically  close  to 
one  another  but  do  not  overlap.  It  is  hard 
to  imagine  that  the  hurricanes  that  fre- 
quently pass  through  the  Caribbean,  as 
well  as  the  homogeneous  island  environ- 
ments, would  not  gradually  cause  these 
two  species  distributions  to  overlap.  Per- 
haps these  abrupt,  disjunct  distributions 
are  a  rare  example  of  competitive  exclu- 
sion in  Metepeira,  which  in  other  species 
is  not  thought  to  be  an  important  factor 
(Wise,  1983). 

Sphecid  wasps  are  predators  on  Mete- 
peira. Locality  labels  indicated  that  M.  pa- 
cifica  has  been  found  in  the  nests  of  Try- 
pargilum  nitidum,  T.  tenoctitlan,  and  T. 
hensoni.  Jimenez  and  Tejas  (1994)  report 
that  M.  crassipes  is  the  most  frequent  prey 
item  in  the  nests  of  Trypargilum  triden- 


tatum.  Colonial  spiders,  such  as  M.  incras- 
sata, are  especially  vulnerable  to  wasps, 
other  spiders,  sarcophagid  flies  (e.g.,  Ar- 
achnidomyia  Undue,  A.  rayorae),  and 
hummingbirds  (Hieber  and  Uetz,  1990; 
Lopez,  1989;  Rayor  and  Uetz,  1990). 

Species  Groups.  Nearctic  Metepeira 
were  divided  into  two  species  groups:  the 
M.  labyrinthea  group  and  the  M.  foxi 
group,  based  on  the  pattern  on  the  ster- 
num and  the  shape  of  the  median  apoph- 
ysis (Levi,  1977).  Baert  (1987)  questioned 
the  taxonomic  usefulness  of  the  M.  foxi 
species  group  (M.  foxi,  M.  grandiosa,  M. 
datona)  because  he  found  that  M.  desen- 
deri  has  both  a  keel  on  the  median  apoph- 
ysis and  a  white  sternal  line  (Figs.  15, 
21) — a  combination  that  is  incoinpatible 
by  Levi's  scheme.  Nonetheless,  the  geni- 
talia of  M.  desenderi  closely  ally  this  spe- 
cies with  the  M.  foxi  group,  so  I  am  re- 
defining the  M.  foxi  group  based  on  purely 
genitalic  characters.  This  is  likely  to  be  a 
basal,  paraphyletic  group  (Fig.  4)  (Piel  and 
Nutt,  1997). 

Seven  additional  species  groups  are  dis- 
tal to  the  M.  foxi.  These  remaining  species 
are  united  by  sharing  a  large  terminal 
apophysis  that  is  sclerotized  and  usually 
studded  with  teeth  or  denticles.  The  M. 
vigilax  group  (M.  vigilax,  M.  cajahamha, 
M.  glomerahilis,  M.  rectangida)  are  united 
by  large  emboli  with  long  scooplike  basal 
embolic  apophyses  (Fig.  60).  Unlike  the 
remaining  species,  the  terminal  apophysis 
in  this  group — albeit  large — does  not  ac- 
tually overhang  or  hide  the  embolus.  In 
addition  to  an  overhanging  terminal 
apophysis,  the  remaining  taxa  are  also 
united  by  a  distal  embolic  apophysis  that 
either  protrudes  (Fig.  76),  curves  off  (Fig. 
185),  or  is  secondarily  lost  (Fig.  264).  The 
M.  labyrinthea  group  (M.  labyrinthea,  M. 
lacandon,  M.  spinipes)  share  a  toothless, 
smooth  keel  on  the  median  apophysis 
(Figs.  67,  69). 

The  M.  nigriventris  group  and  the  M. 
compsa  group  together  share  a  median 
apophysis  with  teeth  on  the  face  of  the 
keel   (Figs.   92,    149).   The  M.    incrassata 


Metepeira  •  Piel 


11 


Map  1 


•  glomerabilis 
+  cajabamba 


Map  3 


Map  2 


grandiosa  alpina 
grandiosa  grandiosa' 


4»^ 


•  vigilax 
+  rectangula 

Map  4 


Maps  1,  2.  Metepeira  fox;  species  group.  1,  M.  datona,  M.  desenderi.  2,  M.  grandiosa  grandiosa.  M.  grandiosa  alpina. 
Maps  3,  4.  Metepeira  vigilax  species  group.  3,  M.  glomerabilis.  M.  cajabamba.  4,  M.  vigilax,  M.  rectangula. 


group,  the  M.  ventura  group,  and  the  M. 
minima  group  all  lack  a  keel  on  the  me- 
dian apophysis  (Figs.  164,  222,  293).  How- 
ever, both  the  M.  compsa  group  and  the 
M.  incrassata  group  have  epigyna  with 
similar  oval  or  round  sclerotized  rims 
(Figs.  151,  166),  so  it  is  likely  that  these 
are  paraphyletic  and  consist  of  species 
leading  up  to  a  major  North  American 
(without  a  keel)  and  South  American  (with 
a  keel)  phylogenetic  split  (Fig.  4). 

The  South  American  branch  includes 
the  M.  compsa  group  (M.  compsa,  M.  ro- 
raima,  M.  gressa)  and,  more  distally,  the 
M.  nigriventris  group  (M.  nigriventris,  M. 
tarapaca,  M.  calamuchita,  M.  galatheae, 
M.  karkii).  This  latter  group  is  united  by  a 


distinctive  and  derived  scape,  which  pro- 
jects out  and  down,  creating  a  noticeable 
arch  and  overhang  (Fig.  86). 

The  remaining  species  all  lack  a  keel  on 
the  median  apophysis,  and  with  one  ex- 
ception {M.  inca),  they  live  exclusively  in 
North  America  (from  the  Caribbean  and 
Panama  to  Nevada).  The  M.  incrassata 
group  (M.  gosoga,  M.  maya,  M.  inca,  M. 
Comanche,  M.  olmec,  M.  atascadero,  M.  ar- 
izonica,  M.  incrassata,  M.  triangularis,  M. 
pimungan)  are  very  likely  paraphyletic. 
The  epigynum  on  each  species  seems  au- 
tapomorphic  and  difficult  to  unite  with  any 
others.  Some  species  (M.  gosoga,  M.  maya, 
M.  inca)  have  a  pointed  or  projecting  distal 
embolic  apophysis  (Fig.  171).  Others  have 


12         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


a  distal  embolic  apophysis  that  curves  off 
sharply  but  does  not  project  forward  (Fig. 
185).  Finally,  others  have  a  distal  embolic 
apophysis  that  curves  off  gently,  almost  to 
the  point  of  hiding  the  existence  of  an 
apophysis  (Fig.  206). 

The  M.  minima  group  (M.  jamaicensis, 
M.  minima,  M.  pacifica,  M.  petatlan)  and 
the  M.  Ventura  group  (M.  uncata,  M.  Ven- 
tura, M.  celestun,  M.  chilapae,  M.  revilla- 
gigedo  new  species,  M.  crassipes)  are  unit- 
ed by  derived  characters:  both  have  thin 
flagellae  arising  from  a  thin  base  on  the 
median  apophysis  (Fig.  264),  and  both 
share  the  secondary  loss  of  the  distal  em- 
bolic apophysis.  The  M.  minima  group  is 
clearly  monophyletic;  its  species  all  have 
their  flagellae  further  set  off  from  the  me- 
dian apophysis  on  a  separate,  narrow  stalk 
(Fig.  286). 

Key  to  Female  Metepeira 

1  Epigynal  openings  strongly  sclerotized, 
round  but  tilted  so  that  they  appear 
oval  from  a  ventral  view  (Figs.  40,  48, 
55)  2 

—  Epigynal   openings   weakly   sclerotized 

(e.g..  Fig.  208),  and  if  round,  they  are 
not  tilted  and  do  not  appear  oval  from 

a  ventral  view  (Fig.  131)   4 

2(1)  Epigynal  openings  shaped  like  the  en- 
trance to  a  snail  shell  (Fig.  55);  His- 

paniola,  Brazil,  Argentina  (Map  4) 

( 7 )  vigilax 

—  Epigynal  openings  tubular  (Figs.  40,  48) 

3 

3(2)        Epigynal   openings   strongly  tilted   (Fig. 

40);  Peru  (Map  3)  (5)  cajabamba 

—  Epigynal  openings  weakly  tilted  (Fig.  48); 

Colombia  to  Brazil  (Map  3)  

(6)  glomerabilis 

4(1)  Epigynal  openings  large  and  gaping,  cre- 
ating large  atria  inside  (Figs.  61,  62); 

Chile  and  Argentina  (Map  4)   

( 8 )  rectangula 

—  Epigynal   openings   not  gaping  and  not 

creating  large  atria  (e.g..  Fig.  69)  5 

5(4)  Weak  posterior  lobes  on  the  epigynum 
create  a  single,  wide  epigynal  depres- 
sion (Figs.  9,  17,  24,  31)  or  epigynum 
with  crescent-shaped  sclerotized  open- 
ings on  either  side  of  a  thin  scape 
(Levi,  1977,  fig.  87).  Dark,  sclerotized 
spheres  below  epigynal  openings  are 
greatly  separated  (Figs.  8,  16)  6 

—  Stronger  posterior  lobes  on  the  epigynum 


create  separate,  smaller  epigynal  de- 
pressions that  are  not  crescent-shaped, 
or  if  crescent-shaped,  the  scape  is  thick 
and  puffy  (e.g..  Figs.  86,  102,  143,  166, 
187,  208).  Dark,  sclerotized  spheres 
below  epigynal  openings  are  closer  to- 
gether (e.g..  Figs.  101.  142)  11 

6(5)        Sternum   with   longitudinal  white   line 
(Fig.  21);  Galapagos  Islands  (Map  1) 
(2)  desenderi 

-  Sternum  entirely  black  or  brown  (Figs. 

13,  28,  35)   7 

7(6)  Abdomen  wider  than  long  (Fig.  12);  Flor- 
ida to  Hispaniola  (Map  1)  (1)  datona 

-  Abdomen  longer  dian  wide  (Figs.  27,  34) 

8 

8(7)  Coxae  as  black  as  sternum  (Levi,  1977, 
fig.    98);    Canada-U.S.    border    (Levi, 

1977,  map  2)  

(Levi,  1977:  212)  grandiosa  palustris 

—  Coxae  yellow  or  orange  and  lighter  than 

sternum  (Figs.  28,  35)  9 

9(8)  Epigynum  with  crescent-shaped  sclero- 
tized openings  on  eitlier  side  of  die 
scape  (Levi,  1977,  fig.  87);  western 
U.S.  and  Canada  (Levi,  1977,  map  2) 
(Levi,  1977:  210)  foxi 

—  Epigynum  with  wide,  transverse  depres- 

sion (Figs.  23,  24,  30,  31)   10 

10(9)     Scape  wide  and  stubby  (Fig.  24);  Baja 
California  north  to  Canada  (Map  2) 
(3)  grandiosa  grandiosa 

-  Scape  triangular  (Fig.  31);  in  mountains 

from  north-central  Mexico  to  Canada 

(Map  2)  (4)  grandiosa  alpina 

11(5)     Scape   diickness   equal   to   greater  than 
width    of   epigynal    depressions    (e.g.. 
Figs.  86,  119,  131,  143,  180,  201,  266)       ' 
12 

—  Scape    narrower   than    epigynal   depres- 

sions, or  epigynal  depressions  in  the 
shape  of  longitudinal  slits  (e.g..  Figs. 

238,  252,  280)  35 

12(11)  Base  of  scape  originates  anteriorly  and 
projects  ventrally  before  cui'ving  pos- 
teriorly. This  projection  creates  an 
overhang  and  a  noticeable  gap  between 
the  scape  and  the  genital  openings 
(e.g..  Figs.  85,  86,  101,  102,  119,  143) 
13 

—  Scape  does  not  create  a  noticeable  gap  or 

overhang  (e.g..  Figs.  77,  78,  130,  131, 

165,  166,  265)  18 

13(12)  Rim  of  epigynal  depressions  slightly 
sclerotized  and  oval-shaped  (Fig.  143); 
northern  Brazil,  French  Guiana,  and 
Colombia  (Map  8)   (17)  roraima 

-  Epigynal  depressions  without  distinct  rim 

(Fig.  123)  or  not  sclerotized  (e.g..  Fig. 

102)  14 

14(13)   Sternum  black,  coxae  mostly  black  (Fig. 


Metepeira  •  Piel 


13 


A  calamuchita 


karkii 


mgnventris   f  (       ^ 


Map  5 


▲  galatheae 
•  tarapaca 


Map  7 


•  spinipes 
■  lacandon 


Map  6 


+  compsa 
•  roraima 
■  gressa 


Map  8 


Maps  5,  7.  Metepeira  nighventris  species  group.  M.  calamuchita,  M.  I<arl<ii,  M.  nigriventris,  M.  galattieae,  M.  tarapaca. 
Map  6.  Metepeira  labyrintfiea  species  group.  M.  spinipes,  M.  lacandon. 
Map  8.  Metepeira  compsa  species  group.  M.  compsa,  M.  roraima,  M.  gressa. 


91),  and  carapace  without  lighter  me- 
dian mark  (Fig.  90);  high  altitudes  in 
Bolivia  and  Peru   near  Lake  Titicaca 

(Map  5)  (11)  nigriventri.s 

Sternum  with  median  white  line  (Fig. 
99),  or  if  sternum  is  black,  then  either 
the  carapace  has  a  median  lighter  ar- 
row-shaped mark  (Fig.  113)  or  the  cox- 
ae are  mostly  yellow  (Figs.  114,  128) 
15 


15(14)  Stenium  brown  to  black  with  parallel 
lines  on  either  side  of  median  white 
line  on  venter:  the  parallel  lines  are 
tliicker  anteriorly  than  posteriorly  (Fig. 
128).  Lower  lip  on  epigynum  thick  and 
bulbous  (Fig.  123);  southern  Argentina 
and  southern  Chile  (Map  5)  ....  (15)  karkii 

—  Sternum  with  median  white  line,   or  if 

sternum  is  entirely  brown  to  black, 
then   parallel   lines   on   either  side   of 


14         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


white  line  on  venter  are  either  absent 
(Fig.  114)  or  equally  thick  anteriorly  as 
posteriorly  (Fig.  99).  Epigynum  with- 
out thickened  lower  lip  (Figs.  94,  102, 

118,  119,  120)  16 

16(15)  Dark,  sclerotized  spheres  in  epigynal 
openings  small  (Figs.  119,  120).  Ante- 
rior lip  of  epigynum  rounded  off, 
sometimes  with  openings  shifted  pos- 
teriorly and  wiinkled  portion  of  the 
scapes  hood  shifted  more  to  the  epi- 
gynum proper  (Fig.  118);  Chile  and 
Argentina  (Map  7)   (14)  galatheae 

—  Dark,    sclerotized    spheres    in    epigynal 

openings  large,  and  anterior  lip  of  epi- 
gynum not  rounded  off.  Openings  al- 
ways located  midway  down  the  epigyn- 
um (Figs.  94,  102)   .._._ 17 

17(16)  General  shape  of  epigynum  is  triangular 
with  posterior  width  greater  than  an- 
terior width.  Scooped-out  depressions 
project  more  posteriorly  than  laterally 
(Fig.  102);  north-central  Argentina 
(Map  5)  (13)  calamiichita 

—  General    shape    of    epigynum    square. 

Scooped-out  depressions  project  more 
laterally  than  posteriorly  (Fig.  94); 
northern    Chile    and    southern    Peru 

(Map  7)  (12)  tarapaca 

18(12)  Epigynal  openings  almond-shaped,  not 
noticeably  sclerotized,  and  at  a  40°  to 
60°  angle  from  axis  of  spider  (Figs.  69, 
78)  19 

—  Epigynal    openings    not    almond-shaped 

(e.g..  Figs.  180,  187,  201),  or  if  al- 
mond-shaped, slightly  sclerotized  and 
at  an  angle  from  spider's  axis  of  80°  to 

100°  (Figs.  151,  159,  166)  22 

19(18)  Almond-shaped  openings  created  by  de- 
pression where  scape  arises  from  epi- 
gynum (Fig.  173);  California,  Arizona, 

northwestern  Mexico  (Map  9)   

(21)  gosoga 

—  Almond-shaped    openings    created    by 

membranes  inside  depressions  and  not 
associated  with  scape  (Figs.  69,  78)  .....  20 
20(19)  Distinct  C-shaped  depression  created 
where  die  scape  arises  from  the  epi- 
gynum (Fig.  78).  Black  marks  inside  al- 
mond-shaped openings  not  cross-eyed 
in  appearance  (Fig.  78).  Black  ster- 
num, usually  with  white  spot  in  center 
(Fig.  83);  mountainous  regions  of  Chia- 
pas, Mexico  (Map  6)   (10)  lacandon 

—  Indistinct  depression  created  where  the 

scape  arises  from  the  epigynum  (Fig. 
69).  Black  marks  inside  almond-shaped 
openings  cross-eyed  in  appearance 
(Fig.  69).  Sternum  black,  with  or  with- 
out a  median  white  line.  If  with  only  a 
portion  of  a  median  white  line  present. 


usually  only  at  the  posterior  end  of  the 
sternum  (Fig.  75).  Never  with  only  one 

white  spot  in  center 21 

21(20)  Because  of  interspecific  variability  and 
polymorphism,  females  of  the  follow- 
ing two  species  are  almost  impossible 
to  separate  reliably  without  molecular 
sequence  data.  Small  ribosomal  sub- 
unit  (12S)  mtDNA  sequence  data  has 
tlie  following  diagnostic  markers.  Base 
14261:  ACGGT;  base  14285:  ATTTT; 
base  14361:  ACTAC;  base  14394: 
CTTAT;  base  14412:  ATTA.  (Base 
numbers  refer  to  homologous  sites  in 
tlie  mitochondrion  of  Drosophila  yak- 
uba,  as  reported  by  Clary  and  Wolsten- 
holme  [1985].)  One  quarter  of  scape 
extends  below  lower  lips  of  epigynum 
(Levi,  1977,  fig.  14);  New  England  to 
Florida  and  west  to  eastern  Texas 
(Levi,  1977,  map  1,  but  not  including 

points  appearing  in  Mexico)  

(Levi,  1977:  196)  labyrinthea 

—  For  12S  mtDNA  sequence  data,  the  fol- 

lowing sequences  are  diagnostic.  Base 
14261:  AGGAT;  base  14285:  ATCTT; 
base  14361:  ACCAC;  base  14394: 
CTAAT;  base  14412:  TTTA.  One  third 
of  scape  extends  below  lower  lips  of 
epigynum  (Levi,  1977,  fig.  21;  Fig.  69); 
Mexico  City  north  to  California  (Map 

6)   (9)  .spinipes 

22(18)  Epigynal  openings  small,  round,  and 
sclerotized  (Fig.  131).  Openings  some- 
times hidden  by  wide  scape  (Fig.  134); 
Puerto  Rico  to  Argentina  and  Chile 
(Map  8)  (16)  compsa 

—  Epigynal  openings  not  small,  round,  and 

sclerotized  23 

23(22)  Rim  of  epigynal  openings  sclerotized  and 
in  an  oval  or  teardrop  shape  (Figs.  151, 
159,  166,  187)  24 

—  Rim  of  epigynal  openings  not  sclerotized 

in  an  oval  or  teardrop  shape  (Figs.  180, 

194,  201,  208)  27 

24(23)  Epigynal  openings  oval,  small,  and  partly 
hidden  by  scape  (Fig.  151);  northern 
Argentina,  Uruguay,  and  southern  Bra- 
zil (Map  8)  (18)  gressa 

—  Epigynal  openings  teardrop-shaped  (Fig. 

187)  or  large  and  oval  but  not  hidden 

by  scape  (Figs.  159,  166)  _. 25 

25(24)  Epigynal  openings  teardrop-shaped  (Fig. 
187);    northeastern    Mexico    to    Texas 

(Levi,  1977,  map  1;  Map  11)  

(23)  Comanche 

—  Epigynal    openings    oval-shaped   (Figs. 

159,  166)  26 

26(25)  Lower  lip  of  epigynum  pointed  (Fig. 
166),  abdomen  white  (Fig.  169);  north- 
ern tip  of  Peru  (Map  11)   (20)  inca 


Metepeira  •  Piel 


15 


+  arizonica  -x 

•  gosoga 

♦  incrassata 
■  pimungan 


Map  9 


pacifica 
petatlan 


Map  10 


+  Comanche 
X  inca 
♦  olmec 


Map  11 


•  uncata 

♦  crassipes 
■  revillagigedo 


Map  12 


Map  13 


■  Ventura 
▲  celestun 
•  chilapae 


jamaicensis 
minima  b;  s 


jcmaya 
+  atascadero 
•  triangularis 
Map  14 


^;:^5y-.^ 


Map  15 


Maps  9,  11,  14.  Metepeira  incrassata  species  group.  9,  M.  arizonica,  M.  gosoga,  M.  incrassata,  M.  pimungan.  ^^,  M.  comanche, 

M.  inca,  M.  olmec.  14,  M.  maya,  M.  atascadero,  M.  triangularis. 

Maps  10,  15.  Metepeira  minima  species  group.  10,  M.  pacifica,  M.  petatlan.  15,  M.  minima,  M.  jamaicensis. 

Maps  12,  13.  Metepeira  ventura  species  group.  12,  M.  uncata,  M.  crassipes,  M.  revillagigedo.  13,  M.  ventura,  M.  celestun,  M. 

chilapae. 


—  Lower  lip  of  epigynum  thickened  but  not 

pointed  (Fig.  159),  abdomen  dark  (Fig. 
162);  southern  Mexico  and  Belize  to 

Costa  Rica  (Map  14)  (19)  maya 

27(23)  Rim  of  epigynal  openings  sclerotized  and 
shaped  hke  a  pair  of  sunglasses  (Fig. 
201);  eastern  Cuba  and  Hispaniola 
(Map  14)  (25)  triangularis 

-  Rim  of  epigynal  openings  not  sclerotized 


or,  if  sclerotized,  nol  in  shape  of  sun- 
glasses (e.g..  Figs.  208,  224) 28 

28(27)  Posterior  epigynal  lobes  converge  behind 
the  scape  so  that  epigynal  depressions 
appear  closed  off  froin  ventral  view 
(Figs.  180,  208,  216,  224)   29 

-  Posterior  epigynal  lobes  end  before  they 

disappear  behind  the  scape  so  that  epi- 
gynal depressions  appear  open  poste- 


16         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


riorly   from   ventral   view    (Figs.    194, 

245,  259,  266)  32 

29(28)  Scape  relatively  long  and  thin,  epigynal 
depressions  large  and  round,  large 
black  spheres  take  up  almost  all  the 
space  in  the  depressions  (Fig.  180); 
southern  Veracruz  to  Panama  (Map  11) 
(22)  olmec 

—  Scape    relatively   short,    fat,    and   fleshy. 

Epigynal  depressions  not  perfectly 
round;  black  spheres  do  not  take  up 
most  of  the   space  in  the  depression 

(Figs.  208,  216,  224)  30 

30(29)  Epigynum  puffy,  scape  so  thick  that  de- 
pressions on  either  side  of  scape  ap- 
pear crescent-shaped  (Fig.  208);  south- 
western U.S.  to  central  Mexico  (Map 
9)   (26)  arizonica 

—  Epigynum  not  puffy,  scape  not  so  fat  that 

depressions    become    crescent-shaped 

(Figs.  216,  224)  31 

31(30)  Very  social.  Epigynal  depressions  large, 
disk-shaped,  with  shiny-smooth  scler- 
otized  inner  surfaces  and  thin  posterior 
lips  (Fig.  224).  Sternum  mostly  black, 
sometimes  with  anterior  white  marks. 
Venter  without  U-shaped  mark  circum- 
scribing median  white  line  posteriorly. 
Coxae  mostly  dark  brown  (Fig.  228); 
mainly  in  southern  Veracruz  (Map  9) 
(28)  incrassata 

—  Mostly    solitary.    Epigynal    depressions 

small,  oval,  with  reduced  shiny-smooth 
sclerotized  surfaces  and  thick  posterior 
lips  (Fig.  216).  Sternum  with  median 
white  line  or  only  white  mark  at  pos- 
terior end.  Venter  with  U-shaped  mark 
circumscribing  median  white  line  pos- 
teriorly. Coxae  mostly  yellow  (Fig. 
221);    central    Mexican   plateau    (Map 

14)  (27)  atascadero 

32(28)  Small  square-shaped  epigynal  depres- 
sions on  either  side  of  scape  (Figs.  259, 
266)  33 

—  Large,   more   rounded  epigyniil   depres- 

sions (Fig.  194)  or  with  straight  to  S- 
shaped  edges  mostly  covered  by  scape 

(Fig.  245)  34 

33(32)  Dark  spheres  inside  epigynal  depressions 
appear  slightly  walleyed.  Scape  rela- 
tively narrower  at  base:  about  the 
width  of  the  depressions  (Fig.  266); 
southeast  and  south-central  Mexico 
(Map  13)  (34)  chilapae 

—  Dark  spheres  inside  epigynal  depressions 

appear  slightly  cross-eyed.  Scape  rela- 
tively wider  at  base:  about  twice  the 
width  of  the  depressions  (Fig.  259); 
northwestern    Mexico    and    California 

(Map  12)  (33)  crassipes 

34(32)  Large,     rounded     epigynal     depressions 


with  large  black  spheres  inside  (Fig. 
194).  Sternum  black  with  posterior 
dewdrop-shaped  mark.  Fair  of  short 
parallel  lines  on  either  side  of  ventral 
median  white  line  (Fig.  198).  Carapace 
with  large  anterior  white  region  (Fig. 
197);  San  Nicolas  Island  off  southern 
California  (Map  9)  (24)  pimungan 

—  Epigynal    depressions    with    straight    to 

slightly  S -shaped  edges,  mostly  hidden 
by  a  wide  triangular  scape  (Fig.  245). 
Sternum  with  wide  median  white  line. 
No  parallel  white  lines  on  either  side 
of  ventral  median  white  line  (Fig.  249). 
Carapace  with  small  anterior  white  re- 
gion   (Fig.    248);    Yucatan    Peninsula 

(Map  13)  (31)  celestun 

35(11)  Epigynal    depressions    wider   than    long 

(Figs.  231,  252)  36 

—  Epigynal   depressions  longer  than  wide 

(e.g..  Figs.  238,  273,  280,  288)  37 

36(35)  Black  comma  shapes  inside  epigynal  de- 
pressions (Fig.  231).  Sternum  with  me- 
dian white  line  (Fig.  235).  Dorsum 
lightly  pigmented  (Fig.  234);  north- 
western Mexico  and  coastal  California 
(Map  13)  (29)  Ventura 

—  Black  S-shaped  marks  inside  epigynal  de- 

pressions (Fig.  252).  Sternum  black 
with  dewdrop  mark  at  posterior  end 
(Fig.  256).  Dorsum  darkly  pigmented 
(Fig.  255);  Guatemala  and  Costa  Rica 

(Map  12)  ___  (32)  uncata 

37(35)  Epigynal  depressions  indistinct  anterior- 
ly. Black  comma-shaped  marks  inside 
depressions  and  covered  by  translucent 
membranes  (Fig.  238);  Isla  Socorro  of 
the  Archipielago  de  Revillagigedo 
(Map  12)  (30)  revillogigedo 

—  Epigynal  depressions  distinct  anteriorly. 

Black  spheres  shifted  laterally  and  lo- 
cated   outside    the    depressions    (e.g.. 

Figs.  273,  280,  295)  38 

38(37)  Epigynal  depressions  slit-shaped  and 
usually  narrower  than  scape  (Figs.  280, 
281);  northwestern  Mexico  and  Yuca- 
tan Peninsula  (Map  15)  (36)  minima 

—  Epigynal  depressions  oval  and  wider  than 

scape  (Figs.  273,  288)  39 

39(38)  Dark  spheres  larger  than  epigynal  open- 
ings (Fig.  273);  west  coastal  Mexico 
(Map  10)  (35)  petatlan 

—  Dark  spheres  smaller  than  epigynal  open- 

ings (Figs.  288,  295)  40 

40(39)  V-shaped  ridge  under  die  scape.  Dark 
spheres  located  behind  the  junction 
where  the  ridge  meets  the  lateral  edge 
of  the  epigynal  depressions  (Fig.  288); 

Honduras  to  Costa  Rica  (Map  10)  

(37)  pacifica 

—  Straight    ridge    under    the    scape.    Dark 


Metepeira  •  Piel 


17 


spheres  located  laterally  and  outside  of 
the  junction  where  the  ridge  meets  the 
lateral  edge  of  the  epigynal  depressions 
(Fig.  295);  Cayman  Islands,  Jamaica, 
and  Haiti  (Map  15)  .—  (38)  jamaicensis 

Key  to  Male  Metepeira 

1  Terminal  apophysis  tliin,  small,  fleshy, 
without  teeth  or  sclerotized  parts  (Figs. 
7,  15,  22,  29) 2 

-  Terminal  apophysis  enlarged,  meaty,  with 

teeth  or  sclerotized  parts   (e.g..   Figs. 

38,  60,  84,  199) 7 

2(1)  Terminal  apophysis  narrow;  embolus 
curled  clockwise  like  the  tip  on  a  cork- 
screw (Figs.  7,  15)   3 

-  Terminal  apophysis  wide;  embolus  tilted 

up  and  L-shaped  (Figs.  22,  29)  4 

3(2)  Curled  embolus  with  a  ridge  on  the  up- 
per surface  and  raised  on  a  pedicel; 
basal  embolic  apophysis  enormous 
(Fig.  15).  Sternum  witli  wide  median 
white  line  (Fig.  21);  Galapagos  Islands 
(Map  1)  (2)  desenderi 

-  Curled  embolus  smooth  on  its  upper  sur- 

face and  not  raised  on  a  pedicel;  basal 
embolic  apophysis  hardly  noticeable 
(Fig.  7).  Sternum  entirely  black  (Fig. 
11);  Florida  to  Hispaniola  (Map  1)  -  -- 

( 1 )  datona 

4(2)  L-shaped  embolus  at  an  acute  (<  90°)  an- 
gle (Levi,  1977,  figs.  91-93);  western 
U.S.  and  Canada  (Levi,  1977,  map  2) 
(Levi,  1977:  210) /oxi 

-  L-shaped   embolus    at   angle    of  90°    or 

greater  (Figs.  22,  29)  5 

5(4)  Median  apophysis  widi  rounded  projec- 
tion on  dorsal  side;  jagged  posterior 
edge  of  keel  (Figs.  22,  29)  6 

-  Median  apophysis  without  projection — 

flat  on  dorsal  side;  rounded  posterior 
edge  of  keel  (Levi,  1977,  fig.  105); 
along  the  Canadian  and  U.S.  border, 
north  to  Nova  Scotia  and  British  Co- 
lumbia, south  to  Maine  and  North  Da- 
kota (Levi,  1977,  map  2)  

(Levi,  1977:  212)  grandiosa  palustris 

6(5)  Lower,  transverse  part  of  L-shaped  em- 
bolus longer  than  vertical  part  (Fig. 
29);    mountains    from    north-central 

Mexico  to  Canada  (Map  2)  

(4)  grandiosa  alpina 

-  Lower,  transverse  part  of  L-shape  em- 

bolus shorter  than  or  equal  to  vertical 
part  (Fig.  22);  Baja  California  north  to 
Canada  (Map  2)  ..  (3)  grandiosa  grandiosa 
7(1)  Terminal  apophysis  does  not  entirely 
overhang  the  embolus.  Embolus  long 
and  robust,  wdth  a  long  and  thin  gap 
created  between  the  embolus  and  the 


basal  ennbolic  apophysis  (Figs.  38,  46, 
53,  60)  8 

-  Terminal  apophysis  often  overhangs  the 

embolus,  covering  it  from  view.  Em- 
bolus not  long  and  robust,  without 
long,  thin  gap  between  embolus  and 
basal  embolic  apophysis  (e.g..  Figs.  67, 

79,  92,  121,  141,  185)  11 

8(7)  Longer  flagellum  as  thick  as  shorter  fla- 
gellum.  Keel  short,  slim,  and  feather- 
shaped.  Embolus  as  wide  as  base  of 
median  apophysis  (Fig.  53);  Hispanio- 
la, Brazil,  Argentina  (Map  4)  ....  (7)  vigilax 

-  Longer  flagellum  thicker  than  shorter  fla- 

gellum. Keel  absent  (Fig.  38)  or  wide 
and  arrowhead-shaped.  Embolus  thin- 
ner  than    base    of  median    apophysis 

(Figs.  38,  46,  60)  9 

9(8)        Keel  absent  or  greatly  reduced  (Fig.  38); 

Pei-u  (Map  3)  (5)  cajabamba 

-  Keel  present  (Figs.  46,  60)   10 

10(9)     Large  embolus  as  long  as  basal  embolic 

apophysis   (Fig.   60).    Normal  embolic 

cap;  Chile  and  Argentina  (Map  4)  

(8)  rectangula 

-  Small  embolus  shorter  than  basal  embol- 

ic apophysis,  often  seen  with  winged 
embolic   cap    (Fig.    46);    Colombia   to 

Brazil  (Map  3)   (6)  glomerabilis 

11(7)  Median  apophysis  with  keel  (e.g.,  Figs. 
92,  141)  

-  Median   apophysis   without   keel    (e.g.. 

Figs.  157,  178,  286);  all  North  Ameri- 
can or  Caribbean  species  except  for  M. 

inca  

12(11)   Keel  without  teeth;   smooth  (e.g..   Figs. 

67,  76);  North  America 13 

-  Keel  with  teeth  on  face;  rough  (e.g..  Figs. 

84,  92,  100);  South  America  15 

13(12)  Distal  embolic  apophysis  sleek,  pointed, 
and  feather-shaped  when  viewed  from 
underside  of  terminal  division  (Fig. 
70);  Mexico  City  north  to  California 
(Map  6)  (9)  spinipes 

-  Distal   embolic  apophysis   spoon-shaped 

(Fig.  6)  or  widened  with  bump  (Fig. 

79)  14 

14(13)  Distal  embolic  apophysis  spoon-shaped 
(Fig.  6).  Sternum  reddish  brown  with 
median  white  line.  New  England  to 
Florida  and  west  to  eastern  Texas 
(Levi,  1977,  map  1,  excluding  points  in 
Mexico)  (Levi,  1977:  196)  labijrinthea 

-  Distal  embolic  apophysis  widened  with 

bump  (Fig.  79).  Sternum  black  with  or 
without  faint  white  mark  in  center 
(Fig.  81);  mountainous  regions  of  Chia- 
pas, Mexico  (Map  6)   (10)  lacandon 

15(12)  Distal  embolic  apophysis  a  simple  exten- 
sion that  projects  forward,  parallel  to 
the    embolus.    Keel    usuallv    rounded 


12 


22 


18         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


(Fig.  129);  Puerto  Rico  to  Argentina 
and  Chile  (Map  8)  (16)  compsa 

—  Distal   embolic   apophysis   raised   up   or 

projected  away  from  the  embolus.  Keel 
usually  pointed  or  jagged  (Figs.  84,  92, 

100,  121,  141,  149)  16 

16(15)  Distal  embolic  apophysis  points  up  and 
away  from  the  embolus.  Keel  usually 
jagged  (Figs.  141,  149) 21 

—  Distal  embolic  apophysis  wide  and  raised 

up  on  boss.  Keel  usually  pointed  (Figs. 
84,  92,  100)   17 

17(16)  Dewlap  extension  under  embolus  (Fig. 
100)  curves  under,  narrowing  the  gap 
between  the  embolus  and  the  basal 
embolic  apophysis  by  one-half  (Fig. 
103);  north-central  Argentina  (Map  5) 

(13)  calarnuchita 

No  such  dewlap  (Figs.  84,  92,  110,  121). 
Gap  between  embolus  and  basal  em- 
bolic apophysis  narrow  by  less  than 
one-half  the  widest  distance  (Figs.  87, 
95,  117,  124)  18 

18(17)  Outside  edge  of  distal  embolic  apophysis 
gently  rounded  when  viewed  from  un- 
derside of  terminal  division  (Fig.  124); 
southern  Argentina  and  southern  Chile 
(Map  5) (15)  karkii 

—  Outside  edge  of  distal  embolic  apophysis 

with  distinct  bump  when  viewed  from 
underside   of  terminal   division   (Figs. 

87,  95,  117)  19 

19(18)  Outside  edge  of  distal  embolic  apophysis 
with  rounded  bump  that  is  sclerotized 
and  has  a  distinct  line  rising  up  from 
embolus  (Fig.  117);  Chile  and  Argen- 
tina (Map  7)  (14)  galatheae 

—  Outside  edge  of  distal  embolic  apophysis 

with  pointed  bump,  less  sclerotized 
than  embolus  proper  and  without  a  dis- 
tinct line  rising  up  from  the  embolus 

(Figs.  87,  95)  20 

20(19)  Thick  neck  joining  embolus  and  basal 
embolic  apophysis.  Bump  on  distal  em- 
bolic apophysis  peeks  out  from  under 
the  terminal  apophysis  (Figs.  92,  95). 
Sternum  usually  with  white  spot  or  me- 
dian   white    line    (Fig.    97);    northern 

Chile  and  southern  Peru  (Map  7)  

(12)  tarapaca 

—  Thin  neck  joining  embolus  and  basal  em- 

bolic apophysis.  Bump  on  distal  em- 
bolic apophysis  does  not  peek  out  from 
under  the  terminal  apophysis  (Figs.  84, 
87).  Sternum  always  black,  general  col- 
oration dark  (Figs.  88,  89);  high  alti- 
tudes in  Bolivia  and  Peru  near  Lake 

Titicaca  (Map  5)  (11)  nigriventris 

21(16)  Distal  embolic  apophysis  thinner  tlian 
embolus  near  die  junction  of  the  two. 
Embolus    tip    is    curved    gendy    (Fig. 


149);  northern  Argentina,  Uruguay, 
and  southern  Brazil  (Map  8)  ...  (18)  gressa 

—  Distal  embolic  apophysis  the  same  size  as 

embolus  near  the  junction  of  the  two. 
Embolus  tip  is  curved  abruptly  (Fig. 
141);  northern  Brazil,  French  Guiana, 

and  Colombia  (Map  8)   (17)  roraima 

22(11)  Distal    embolic    apophysis    a    projecting 

bump  (Figs.  157,  164,  171)  23 

—  Distal    embolic    apophysis    curved    off 

(Figs.  178,  185,  199,  214),  rounded 
(Figs.  192,  206),  or  absent  (Figs.  264, 

278,  293)  25 

2.3(22)  Distal  embolic  apophysis  raised  up  from 
the  embolus  and  pointed  (Fig.  171); 
California,  Arizona,  northwestern  Mex- 
ico (Map  9)  (21)  gosoga 

—  Distal  embolic  apophysis  not  raised  up, 

but  projecting  forward  and  rounded  off 

(Figs.  157,  164)  24 

24(23)  Main  flagellum  on  median  apophysis  thin 
and  initially  as  thick  as  base.  Embolus 
and  embolic  cap  shortened  (Fig.  164); 
northern  tip  of  Peru  (Map  11)  ...  (20)  inca 

—  Main    flagellum    on    median   apophysis 

thick,  but  initially  thinner  than  base. 
Embolus  and  embolic  cap  elongated 
(Fig.  157);  southern  Mexico  and  Belize 

to  Costa  Rica  (Map  14)  (19)  mnija 

25(22)  Embolus  shaped  like  the  nib  on  a  foun- 
tain pen,  with  bump  near  tip  on  op- 
posite side  of  distal  embolic  apophysis 
(Figs.  214,  217);  central  Mexican  pla- 
teau (Map  14)  (27)  atascadero 

—  Embolus   witliout   such   bump   near  tip 

(e.g..  Fig.  222) 26 

26(25)  Distal  embolic  apophysis  abruptly  ends 
in  sharp  curve  and  flagellae  not  set  off 
on  a  narrow  stalk  (Figs.  178,  185,  199, 
222)  27 

—  Distal  embolic  apophysis  gently  cui-ved 

off  (Figs.  229,  236,  243),  rounded 
(Figs.   192,  206),  or  absent  (e.g..  Fig. 

293)  30 

27(26)  Flagellae  veiy  thin  and  equal  in  length. 
Embolus  long  and  arching,  with  wide 
but  short  embolus  cap.  Overhanging 
terminal  apophysis  covers  only  a  distal 
portion  of  the  sclerotized  part  of  the 
embolus  (Fig.  199);  eastern  Cuba  and 
Hispaniola  (Map  14)  (25)  triangularis 

—  Flagellae  normal  in  thickness  and  usually 

of  different  lengths.  Terminal  apophy- 
sis centered  above  the  entire  sclero- 
tized  portion    of  the    embolus    (Figs. 

178,  185,  222)  28 

28(27)  Not  known  to  be  very  social.  Height  of 
embolus  plus  distal  embolic  apophysis 
just  before  the  latter  curves  off  sharply 
is  equal  to  or  greater  than  length  of 
embolus  tip  distal  to  this  point  (Figs. 


Metepeira  •  Piel 


19 


178,  185).  Sternum  with  median  white 
markings  (Figs.  182,  189)  29 

-  Highly  social  species.  Height  of  embolus 

plus  distal  embolic  apophysis  just  be- 
fore die  latter  curves  off  shai^ply  is  less 
than  the  length  of  the  embolus  tip  dis- 
tal to  this  point  (Fig.  222).  Sternum 
mostly  black;  sometimes  with  anterior 
white  marks.  Coxae  mostly  dark  brown 
(Fig.  226);  mainly  in  southern  Vera- 
cruz, Mexico  (Map  9) (28)  incrassata 

29(28)  Darker,  sclerotized  portion  of  the  embo- 
lus does  not  extend  over  the  hump  of 
the  distal  embolic  apophysis.  Base  of 
embolus  narrower  than  widest  part  of 
the  first  flagellum  (Fig.  178);  southern 
Veracruz,  Mexico  to  Panama  (Map  11) 
(22)  olmec 

-  Darker,  sclerotized  portion  of  the  embo- 

lus extends  over  the  hump  of  the  distal 
embolic  apophysis.  Width  of  embolus 
base  the  same  or  greater  than  widest 
part  of  the  first  flagellum  (Fig.  185); 
northeastern    Mexico   to   Texas    (Levi, 

1977,  map  1;  Map  11)   (23)  comanche 

30(26)   Distal  emboUc  apophysis  rounded  off  to 

form  convex  curve  (Figs.  192,  206)  31 

-  Distal  embolic  apophysis  gently  falls  off 

to  form  concave  shape  (Figs.  229,  236, 

243)  or  absent  (e.g..  Fig.  293)  32 

31(30)  Embolus  S-shaped  (Fig.  192).  Sternum 
black  with  posterior  white  mark.  Ven- 
ter without  white  anchor  shape  inark 
(Fig.    196);    San    Nicolas    Island    off 

southern  California  (Map  9)  

(24)  pimiingan 

-  Embolus     convex    on    upper    surface, 

straight  on  lower  surface  (Fig.  206). 
Sternum  black  with  median  white  line. 
Venter  with  faint  white  anchor-shaped 
mark  posterior  to  median  white  line 
(Fig.  210);  southwestern  U.S.  to  cen- 
tral Mexico  (Map  9)   (26)  arizonica 

32(30)  Flagellae  on  median  apophysis  set  off  on 
separate,  narrow,  stalk  (e.g..  Figs.  278, 
286)  38 

-  Flagellae  on  median  apophysis  not  set  off 

on  a  separate,  narrow,  stalk  (e.g..  Figs. 

229,  264)  33 

33(32)  Larger  flagellum  twice  as  wide  as  smaller 
flagellum  (Fig.  236);  Isla  Socorro  of  the 
Archipielago  de  Revillagigedo  (Map 
12)  (30)  revillagigedo 

-  Larger  flagellum  less  than  twice  as  thick 

as   smaller  flagellum   (e.g..   Figs.   229, 

243)  34 

34(33)  Flagellae  on  median  apophysis  relatively 

thin  (Figs.  250,  264)  35 

-  Flagellae   on   median   apophysis   thicker 

(Figs.  229,  243,  257)  36 

35(34)   Embolus  with  sharp  bend  near  the  tip 


(Fig.  264).  Sternum  with  median  white 
line  (Fig.  268);  southeast  and  south- 
central  Mexico  (Map  13)  (34)  chilapae 

—  Embolus  with  gentle  bend  further  from 

the  tip  (Fig.  250).  Sternum  mosdy 
black  with  small  white  mark  at  poste- 
rior end  of  sternum  (Fig.  254);  Gua- 
temala and  Costa  Rica  (Map  12)  

(32)  uncata 

36(34)  Embolus  with  sharp  bend  near  the  tip 
(Fig.  229);  northwestern  Mexico  and 
coastal  California  (Map  13)  ...  (29)  ventura 

—  Embolus  with  gentle  bend  further  from 

the  tip  (Figs.  243,  257)  37 

37(36)  Sclerotized  portion  of  embolus  about  as 
long  as  the  longer  flagellum  (Fig.  243); 

Yucatan  Peninsula  (Map  13)  

(31)  celestun 

—  Sclerotized  portion   of  embolus   shorter 

than  the  longer  flagellum  (Fig.  257); 
northwestern    Mexico    and    California 

(Map  12)  (33)  crassipes 

38(32)  Embolus  thick  and  tapering  to  a  point 

(Figs.  271,  278)  39 

—  Embolus  thin  and  needlelike  (Figs.  286, 

293)  40 

39(38)  Longer  flagellum  greater  than  half  the 
length  of  the  cymbium  (Fig.  271);  west 
coastal  Mexico  (Map  10)  (35)  petatlan 

—  Longer  flagellum  less  than  half  the  length 

of  the  cymbium  (Fig.  278);  northwest- 
ern   Mexico    and   Yucatan   Peninsula 

(Map  15)  (36)  minima 

40(38)  Flageflae  set  off  on  long,  thin  stalk  (Fig. 
293);  Cayman  Islands,  Jamaica,  and 
Haiti  (Map  15)  (38)  jamaicensis 

—  Flagellae  set  off  on  short,  diicker  stalk 

(Fig.  286);  Honduras  to  Costa  Rica 
(Map  10)  (37)  pacifica 

Metepeira  foxi  Group 

The  M.  foxi  species  group  {sensu  Levi, 
1977)  and  M.  desenderi  share  veiy  similar 
genitaha,  especially  among  males.  Conse- 
quently, I  am  expanding  the  M.  foxi  group 
to  include  M.  desenderi,  but  with  the  ex- 
clusion of  the  black  sternum  as  a  diagnos- 
tic character.  The  M.  foxi  group  {sensu 
lato)  includes  males  with  an  embolus  that 
lacks  a  distal  apophysis.  The  embolus  curls 
almost  180°  clockwise  around  a  reduced  or 
fleshy  terminal  apophysis  (Figs.  7,  15,  22, 
29).  In  other  species  groups  the  embolus 
is  straighter  and  does  not  curl  (e.g..  Fig. 
60),  and  the  terminal  apophysis  is  large, 
overhanging  the  embolus,  and  often  scler- 
otized with  teeth  (e.g..  Fig.  121).  Epigynal 


20 


Bulletin  Museum  of  Comparative  Zoologtj,  Vol.  157,  No.  1 


features  uniting  females  in  the  M.  foxi  spe- 
cies group  are  harder  to  discern.  Generally 
speaking,  the  epigynum  has  a  stubby, 
shorter,  often  triangular  scape  and  weaker 
posterior  lobes  that  permit  a  wider  view 
into  the  epigynal  openings  (compare  Figs. 
9,  17,  24,  31  with  Figs.  143,  252). 

1 .  Metepeira  datona 
Chamberlin  and  Ivie 
Figures  7-13,  312;  Map  1 

Metepeira  datona  Chamberlin  and  Ivie,  1942:  68,  fig. 
196,  ? .  Female  holotype  from  Daytona  Beach, 
Florida,  USA,  in  the  AMNH,  examined.  Levi, 
1977:  208-210,  figs.  78-86,  S,  9.  Brignofi,  1983: 
275, 

Metepeira  inenna  Bryant,  1945:  378.  Female  holo- 
type from  Cap  Haitian,  Haiti  in  the  MCZ,  exam- 
ined. First  synonymized  by  Levi,  1977:  208—210. 

Description.  Female  from  Morant 
Point,  Mammee  Bay,  Jamaica.  Carapace 
light  around  eyes  with  short  lateral  poste- 
rior extensions,  sometimes  with  thin  lon- 
gitudinal white  line  (Fig.  12).  Legs  ringed. 
Femur  I  with  row  of  two  macrosetae  on 
anterior  side;  three  light  setae  on  anter- 
oventral  side.  Anterior  half  of  dorsal  foli- 
um white,  posterior  half  black,  margined 
by  white;  two  halves  separated  by  trans- 
verse white  line.  Abdomen  widest  in  cen- 
ter (Fig.  12).  Venter  with  a  longitudinal 
white  line.  Pair  of  white  spots  on  either 
side  of  spiracle  (Fig.  13).  Sternum  usually 
entirely  black,  though  soinetimes  with  thin 
anterior  and  posterior  marks,  suggestive  of 
median  white  line  (Fig.  13).  Ratio  of  eye 
diameters:  posterior  medians  and  anterior 
medians  1.0,  anterior  laterals  1.3,  posterior 
laterals  1.1.  Anterior  median  eyes  separat- 
ed by  1.4  diameters,  posterior  median  eyes 
by  0.9,  anterior  median  eyes  separated 
from  anterior  laterals  by  1.6  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.3  their  diameters.  Total  length  4.2 
mm.  Carapace  1.7  mm  long,  1.4  wide. 
First  femur  2  mm,  patella  and  tibia  2.4, 
metatarsus  1.5,  tarsus  0.8.  Second  patella 
and  tibia  1.9  inm,  third  1.1,  fourth  1.6. 

Male  from  Morant  Point,  Mammee  Bay, 
Jamaica.  Carapace  yellowish  brown,  light 


around  eyes.  Median  white  mark  anterior 
to  thoracic  furrow  (Fig.  10).  Legs  same 
color  as  carapace,  lightly  ringed.  Femur  I 
with  row  of  three  macrosetae  on  anterior 
side,  two  on  anteroventral  side.  Center  of 
dorsum  with  transverse  white  line;  poste- 
rior half  darker  than  anterior  half;  thin  me- 
dian black  line;  margin  of  folium  white 
(Fig.  10).  Venter,  sternum  as  in  female 
(Fig.  11).  Ratio  of  eye  diameters:  posterior 
medians  and  anterior  medians  1.0,  anterior 
laterals  1.3,  posterior  laterals  1.2.  Anterior 
median  eyes  separated  by  1.8  diameters, 
posterior  median  eyes  by  1.1,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  1.5  diameters  of  anterior  lateral  eyes, 
lateral  eyes  separated  by  0.5  their  diame- 
ters. Total  length  3  mm.  Carapace  1.4  mm 
long,  1.1  wide.  First  femur  2.7  mm,  patella 
and  tibia  2.7,  metatarsus  1.9,  tarsus  0.8. 
Second  patella  and  tibia  2.1  mm,  third  1.0, 
fourth  1.3. 

Diagnosis.  The  dorsal  folium  and  ab- 
dominal shape  is  distinctive  in  M.  datona. 
Typically,  the  females  abdomen  is  widest 
in  the  middle,  forming  a  rhomboid  in 
shape  (Fig.  12),  and  both  sexes  have  a 
transverse  white  line  dividing  the  dorsal 
folium,  with  wide  black  markings  on  the 
posterior  half  (Figs.  10,  12).  A  ventral  view 
of  the  epigynum  in  M.  datona  reveals  a 
ridge  under  the  scape  that  almost  forms  a 
straight  line  and  which  cui'ves  up  at  the 
ends  (Fig.  9)  instead  of  a  V-shape  (Figs. 
17,  24,  31).  The  openings  to  the  epigynum 
consist  of  small  slits  that  flank  a  wide  de- 
pression (Fig.  9),  compared  to  larger 
openings  that  are  relatively  closer  together 
(Figs.  17,  24,  31).  The  smooth  embolus  on 
M.  datona  is  somewhat  more  compressed 
as  it  curls  clockwise  like  the  tip  on  a  cork- 
screw, in  contrast  to  the  slightly  ridged 
embolus  supported  on  a  stalk  (Fig.  15)  or 
the  strongly  ridged  and  tilted  embolus 
(Figs.  22,  29).  Like  M.  desenderi  (Fig.  15) 
but  unlike  M.  grandiosa  grandiosa  (Fig. 
22)  and  M.  grandiosa  alpina  (Fig.  29),  tlie 
posteroventral  edge  of  the  keel  on  the  me- 
dian apophysis  is  rounded,  as  opposed  to 
pointed  (Fig.  7).  Finally,  M.  datona  has  the 


Metepeira  •  Piel 


21 


thinnest  and  fleshiest  terminal  apophysis 
in  the  genus  (Fig.  7). 

Variation.  Average  body  length  of  15  fe- 
males examined  3.8  mm,  range  2.9  to  4.8 
mm.  Average  body  length  of  seven  males 
examined  2.6  mm,  range  1.8  to  3  mm. 

Natural  History.  Mature  adults  have 
been  collected  throughout  the  year  (Fig. 
312).  According  to  locality  labels,  M.  da- 
tona  are  found  near  the  beach  on  cactus, 
palm,  low  scrub,  mangrove,  and  bamboo. 
Females  are  known  to  wrap  their  egg  sacs 
in  dead  leaves. 

Distribution.  At  sea  level  in  Florida,  Ba- 
hamas, British  West  Indies,  and  the  Do- 
minican Republic  (Levi,  1977,  map  2;  Map 
1). 

Records  Examined.  BAHAMAS  Abaco  Cays:  Aba- 
co,  26°29'N,  77°5'W,  2.xii.l964  (W.  B.  Peck,  MCZ); 
Allons  Cay,  26°59'N,  77°40'W,  9.V.1953  (E.  B.  Hay- 
den,  AMNH);  Hopetown,  Elbow  Cay,  26°33'N, 
76°57'W,  5.V.1953  (E.  Hayden,  AMNH);  New  Plym- 
outh, Green  Turtle  Cay,  26°50'N,  77°23'W,  7.V.1953 
(G.  Rabb,  AMNH);  Whale  Cay  26°43'N,  77°14'W, 
12.i.l964  (W.  B.  Peck,  CAS).  Acklins  Id.:  Atwood's 
Harbor,  22°13'N,  74°18'W,  15.ix.l958  (A.  W.  Scott  Jr., 
MCZ);  Salina  Point,  22°13'N,  74°18'W,  15.viii.l958 
(R.  Robertson  &  A.  W.  Scott  Jr,  MCZ).  Andros:  Fresh 
Creek,  24°26'N,  77°57'W,  23.iv.1953  (L.  Giovannoh, 
AMNH);  Mangrove  Cay,  24°15'N,  77°39'W, 
26. iv.  1953  (E.  Hayden,  AMNH);  Nicolls  Town, 
25°8'N,  78°0'W,  14.iii.l967  (A.  M.  Nadler,  AMNH). 
Berry  Islands:  Frazier's  Hog  Cay,  25°24'N,  77°50'W, 
29.iv.1953  (E.  Hayden,  AMNH);  Little  Harbor  Cay, 
25°34'N,  77°43'W,  l.vl953  (Hayden  &  Giovannoh, 
AMNH).  Crooked  Island:  North  shore  of  Cripple 
Hill,  22°49'N,  74°16'W,  15.ix.l958  (A.  W.  Scott  Jr, 
MCZ);  NW  end,  Gordon  (=  Gun)  Bluff,  22°50'N, 
74°20'W,  15.viii.l958  (R.  Robinson  &  A.  W.  Scott  Jr, 
MCZ).  Crooked  Island  Group:  Long  Cay  22°37'N, 
74°20'W,  7.iii.l953  (Hayden  &  Giovannoh,  AMNH). 
East  Plana  Cay:  22°37'N,  73°.33'W,  4.iii.l953  (E. 
Hayden,  AMNH).  Exurnas:  Musha  Cay  23°50'N, 
76°15'W,  29.xii.1985  (A.  Boutard,  MCZ);  Warderick 
Wells  Cay  24°22'N,  76°36'W,  9.i.l953  (L.  Giovannoh, 
AMNH),  ll.i.l953  (Hayden  &  Giovannoh,  AMNH). 
Grand  Bahamas  Island:  Dundee  Bay,  26°30'N, 
79°15'W  [?],  25.xii.1965  (L.  Pinter,  MCZ);  near  Fre 
port,  pine-palmetto,  26°34'N,  78°27'W  [?], 
25.vii.1965  (L.  Pinter,  MCZ).  Long  Island:  Clarence 
Town,  23°6'N,  74°59'W,  10.iii.l953  (L.  Giovannoh, 
AMNH);  Deadman's  Cay,  23°14'N,  75°14'W, 
ll.iii.l953  (E.  Hayden,  AMNH).  New  Providence  Is- 
land: 7  mi  W  of  Nassau,  25°5'N,  77°28'W,  4.i.l953 
(Hayden  &  Giovannoli,  AMNH).  North  Bimini: 
25°44'N,  79°15'W,  25.i.l950  (C.  M.  Bogert,  AMNH), 


6.vi.l950  (M.  Gazier  &  F.  Rindge,  AMNH),  15.V.1951 
(W.  J.  Gertsch  &  M.  Gazier,  AMNH),  15.vi.l951  (M. 
Gazier,  P  &  C.  Vaurie,  AMNH),  15.vii.l951  (C.  &  P 
Vaurie,  AMNH),  13.xii.l952  (A.  M.  Nadler,  AMNH), 
28.xi. 19.59  (A.  M.  Nadler,  AMNH).  Rmn  Cay:  near 
Port  Nelson,  2.3°38'N,  74°50'W,  16.ih.l953  (Hayden 
&  Giovannoli,  AMNH).  South  Bimini:  25°42'N, 
79°17'W,  12.vi.l950  (M.  Gazier  &  F  Rindge, 
AMNH),  22.vi.1950  (M.  Gazier  &  F.  Rindge, 
AMNH),  15.V.1951  (W.  J.  Gertsch  &  M.  Gazier, 
AMNH),  15.vii.l951  (C.  &  P  Vaurie,  AMNH), 
4.vLii.l951  (C.  &  P  Vaurie,  AMNH),  4.xii.l952  (A.  M. 
Nadler,  AMNH),  25.ih.1953  (A.  M.  Nadler,  AMNH); 
Gun  Cay  25°35'N,  79°20'W,  15.vi.l951  (AMNH). 
BRITISH  WEST  INDIES  Caicos  Islands:  Long  Cay, 
21°28'N,  71°33'W  10.ii.l953  (E.  Hayden,  AMNH); 
South  Caicos,  from  webs  in  upper  beach  zone, 
21°31'N,  71°30'W,  3.iv.l973  (D.  W  Buden,  MCZ); 
West  Caicos,  21°39'N,  72°28'W,  4.U.1953  (Hayden  & 
Giovannoh,  AMNH),  5.ii.l953  (Hayden,  Rabb,  & 
Giovannoli,  AMNH).  Grand  Cayman  Island: 
19°20'N,  81°10'W,  15.ii.l960  (R.  A.  Levidn,  MCZ). 
CUBA  Oriente:  Banes,  20°58'N,  75°43'W,  2.viii.l955 
(A.  F.  Archer,  AMNH);  Ensenada  Nispero,  Giudamar, 
19°58'N,  75°52'W,  9.xi.l945  (P  Alayo,  AMNH);  Jur- 
agua,  19°56'N,  75°40'W,  l.x.1955  (P  Alayo,  AMNH); 
Santa  Fe  [?],  20°22'N,  75°53'W  (A.  F  Archer, 
AMNH).  DOMINICAN  REPUBLIC  Barahona:  Pla- 
ya  Los  Patos,  17°58'N,  71°11'W,  31.viii.l976  (J.  A. 
Ottenwalder,  MNSD).  La  Altagracia:  Punta  Cana, 
Isla  Saona,  18°8'N,  68°34'W  (Felix  E.  Del  Monte  & 
K.  Guerrero,  MNSD).  HAITI  Dept.  du  Nord:  Cap- 
Haitien,  19°46'N,  72°13'W,  15.iii.l934  (E.  Bryant, 
Utawana  Exp.,  MCZ).  JAMAICA  Portland:  between 
Boston  and  Blue  Hole,  18°6'N,  76°37'W,  29.vii.1955 
(A.  F  Archer,  AMNH).  Saint  Andrews:  Hope  Gar- 
dens, Gordontown,  18°2'N,  76°45'W,  27.vii.1955  (A. 
F  Archer,  AMNH).  Surrey:  Morant  Point,  Mammee 
Bay,  17°53'N,  76°19'W,  14.X.1957  (A.  M.  Chickering, 
MCZ);  Palisadoes,  17°56'N.  76°46'W,  ll.xi.l967  (A. 
M.  Chickering,  MCZ);  Palisadoes  Area,  17°56'N, 
76°46'W,  l.xi.l957  (A.  M.  Chickering,  MCZ).  U.S. 
VIRGIN  ISLANDS  Saint  Thomas:  Morant  Point, 
17°55'N,  76°10'W,  25.vii.1985  (G.  B.  Edwards, 
FSCA). 

2.  Metepeira  desenderi  Baert 
Figures  14-21,  307;  Map  1 

Epeira  labyrinthea:— Banks,  1902:  60.  Banks,  1924: 
97.  Misidentification. 

Metepeira  sp.: — Roth  and  Craig,  1970:  116. 

Metepeira  desenderi  Baert,  1^87:  145,  figs.  16-21,  S , 
9 .  Male  holotype  from  Isla  Pinzon,  Galapagos,  Ec- 
uador, in  the  IRSNB.  Platnick,  1989:  341. 

Note.  Holotype  not  examined  because  the  figures 
in  Baert  (1987)  are  clear  and  because  this  is  the 
only  Metepeira  species  found  on  the  Galapagos. 

Description.  Female  from  east  slope  of 
Isla  Santa  Cruz,  Galapagos  Islands,  Ecua- 


22         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


dor.  Carapace  yellowish  brown,  white 
around  eyes,  lighter  median  line  (Fig.  20). 
Legs  yellowish  white,  slightly  darker  an- 
nulations  on  distal  ends  of  articles.  Femur 
I  with  row  of  four  macrosetae  on  anterior 
side;  two  to  four  on  anteroventral  side. 
Dorsal  fleur-de-lis  pattern  broken  into  four 
white  patches  with  anterior  pair  often  larg- 
er than  in  most  species.  Posterior  pair 
straighter  than  usual,  forming  a  cross  in 
center  of  folium  (Fig.  20).  Venter  olive- 
brown  with  median  white  line  surrounded 
by  white  U-shaped  marking  (Fig.  21).  Pair 
of  white  spots  on  either  side  of  spiracle. 
Sternum  brownish  black  with  wide,  white 
line  widening  anteriorly  (Fig.  21).  Ratio  of 
eye  diameters:  posterior  medians  and  an- 
terior medians  1.0,  anterior  laterals  1.3, 
posterior  laterals  1.3.  Anterior  median 
eyes  separated  by  1.5  diameters,  posterior 
median  eyes  by  0.9,  anterior  median  eyes 
separated  from  anterior  laterals  by  2.3  di- 
ameters of  anterior  lateral  eyes,  lateral  eyes 
separated  by  0.2  their  diameters.  Total 
lengdi  5.5  mm.  Carapace  2.3  mm  long,  1.7 
wide.  First  femur  2.7  mm,  patella  and  tibia 
3,  metatarsus  2.2,  tarsus  0.8.  Second  pateUa 
and  tibia  2.4  mm,  third  1.4,  fourth  2. 

Male  from  same  locality  as  female.  Car- 
apace yellowish  brown  with  wide  white 
median  mark  (Fig.  18).  Legs  ringed  like 
female.  Femur  I  with  row  of  four  macro- 
setae  on  anterior  side;  four  to  seven  on  an- 
teroventral side.  Dorsum  and  venter  as  in 
female,  though  median  white  line  on  ster- 
num sometimes  broken  (Figs.  18,  19).  Ra- 
tio of  eye  diameters:  posterior  medians 
and  anterior  medians  0.9,  anterior  laterals 
1.2,  posterior  laterals  1.1.  Anterior  median 
eyes  separated  by  2.2  diameters,  posterior 
median  eyes  by  1.4,  anterior  median  eyes 
separated  from  anterior  laterals  by  2.3  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.5  their  diameters.  To- 
tal length  3.2  mm.  Carapace  1.6  mm  long, 
1.3  wide.  First  femur  2.6  nam,  patella  and 
tibia  2.6,  metatarsus  2.3,  tarsus  0.8.  Sec- 
ond patella  and  tibia  2.2  mm,  third  1.0, 
fourth  1.6. 

Diagnosis.   Within  the  M.  foxi  species 


0  40         80        120       160 

Collection  day  (since  Jan  1 ) 


200 


Figure  14.  The  length  of  mature  M.  desendeh  collected  on 

specific  days  of  the  year  over  a  period  of  84  years.  There  is  a 

trend  from  smaller  spiders  early  in  the  season  to  large  spiders 

later  in  the  season. 

Scale  of  abscissa:  -80  =  October  12  of  the  previous  year;  40 

=  February  9;  160  =  June  9. 

Symbols:  Females  (o),  stippled  regression  line  (t^  =  40%); 

males  (•),  solid  regression  line  (i^  =  42%). 


group,  the  female  abdomen  shape  of  M. 
desenderi  is  closest  to  M.  datona,  though 
not  quite  as  rhomboid  (compare  Fig.  20 
with  Figs.  12,  27,  34).  The  scape  on  M. 
desenderi  is  narrower  and  less  triangular 
than  other  M.  foxi  species,  and  the  de- 
pression under  the  scape  forms  a  distinct 
U-shaped  smile  that  is  not  seen  in  the  oth- 
ers (compare  Fig.  17  with  Figs.  9,  24,  31). 
Like  M.  datona  (Fig.  7)  but  unUke  M. 
grandiosa  grandiosa  (Fig.  22)  or  M.  gran- 
diosa  alpina  (Fig.  29),  the  posteroventral 
edge  of  the  keel  on  the  median  apophysis 
is  rounded  as  opposed  to  pointed  (Fig.  15). 
Similar  to  M.  datona  (Fig.  7),  the  embolus 
of  M.  desenderi  is  curved  like  a  slightly 
flattened  corkscrew.  Unlike  M.  datona,  the 
embolus  seems  to  be  supported  by  a  ped- 
icel off  a  large  basal  embolic  apophysis 
(Fig.  15).  In  contrast,  a  pedicel  is  not  ev- 
ident in  M.  datona,  and  the  basal  embolic 
apophysis  is  barely  visible  (Fig.  7). 

Variation.  Average  body  length  of  45  fe- 
males exaiTiined  5.8  mm,  range  3.8  to  8.5 
mm.  Average  body  length  of  13  males  ex- 
amined 4.9  mm,  range  2.9  to  6.5  mm. 

Natural  History.  Notes  on  the  collection 
labels  of  Y.  Lubin  and  R.  Silberglied  in- 
dicate that  M.  desenderi  is  active  at  night 


Metepeira  •  Piel 


23 


and  that  the  retreat  is  composed  of  Opun- 
tia  bark  and  croton  leaves.  According  to 
Baert  (1987),  M.  desenderi  is  found  in 
large  numbers  in  arid  ecological  zones  on 
all  islands.  Although  some  mature  speci- 
mens have  been  collected  in  November 
and  August,  most  specimens  are  found  be- 
tween Januaiy  and  June  (Fig.  307),  cor- 
responding to  the  "warm  and  wet"  season 
in  the  Galapagos  (van  der  Werff,  1978). 
The  size  of  mature  M.  desenderi  speci- 
mens appears  to  correlate  with  the  collec- 
tion date:  female  spiders  taken  in  August 
are,  on  average,  80%  larger  than  those  tak- 
en in  November  (Fig.  14).  This  trend  may 
indicate  that  M.  desenderi  can  vary  the 
number  of  instars  before  maturity.  Alter- 
natively, the  intermolt  growth  may  be 
greater  for  spiders  with  their  antepenulti- 
mate or  penultimate  instars  occurring  dur- 
ing the  warmer  and  wetter  season. 

Distribution.  Endemic  to  the  Galapagos 
Islands  (Map  1). 

Records  Examined.  ECUADOR  Galapagos:  Albe- 
marle, Tagus  Cove,  0°16'S  91°22'W,  23.1.1899 
(AMNH),  8.ii.l899  (AMNH),  21.iii.l899  (AMNH), 
23.iii.1899  (AMNH);  Archipielago  de  Galapagos, 
0°0'N,  90°30'W  (Williams  Exped.,  1923,  MCZ);  Ba- 
hia  Conway,  Indefatigable  Island,  0°33'S,  90°32'W, 
17.iii.l935  (Exline-Peck,  CAS);  Barrington  Island, 
0°49'S,  90°4'W,  l.viii.l929  (H.  H.  Cleaves,  CAS); 
Campion,  nr.  Floreana  (Santa  Maria),  1°15'S, 
90°27'W,  l.vi.l981  (Y.  Lubin,  MCZ);  Charles  Island, 
1°17'S,  90°26'W,  10.V1899  (AMNH);  Indefatigable 
Island,  0°38'S,  90°23'W,  27.iv.1899  (AMNH), 
18.vi.l929  (Pinchot  South  Sea  Exp,  USNM);  Isla 
Abingdon,  0°35'N,  90°44'W,  25.vi.1899  (AMNH);  Isla 
Albemarle,  0°30'S,  91°4'W,  13. i. 1899  (AMNH), 
20.ii.l899  (AMNH),  20.iii.l899  (AMNH);  Isla  Bin- 
dloe,  0°19'N,  90°29'W,  20.vi.l899  (AMNH);  Isla 
Hood,  1°23'S,  89°39'W,  18.V.1899  (AMNH), 
26.V1899  (AMNH);  Isla  Pinta,  S  Coast,  0°35'N, 
90°44'W,  25.V1964  (D.  Q.  Cavagnaro,  CAS);  Isla  Pla- 
za, 0°35'S,  90°9'W,  7.iii.l970  (R.  Silberglied,  MCZ), 
26.xi.1973  (Y.  Lubin,  MCZ);  Isla  Santa  Cruz,  Acade- 
my Bay,  0°44'30"S,  90°17'30"W,  13.iii.l970  (R.  Sil- 
berglied, MCZ);  Isla  Santa  Cruz,  E  slope,  0°38'S, 
90°23'W,  16.ivl964  (D.  Q.  Cavagnaro,  CAS);  Isla 
Santa  Cruz,  Estacion  Cientifica  Charles  Darwin, 
0°44'S,  90°18'W,  24.i.l964  (D.  Q.  Cavagnaro  &  R.  O. 
Schuster,  CAS),  12.ii.l964  (Cavagnero  &  Schuster, 
CAS),  3.xi.l973  (Y.  Lubin,  MCZ),  24.xi.1973  (Y  Lu- 
bin, MCZ),  25.xi.1973  (Y.  Lubin,  MCZ);  Isla  Santa 
Fe,  S  coast,  0°50'S,  90°4'W,  30.i.l983  (Y.  Lubin, 
MCZ);  James,  0°16'S,  90°42'W,  22.ivl899  (AMNH); 


Narborough  Island,  0°25'S,  91°30'W,  28.iii.1899 
(AMNH);  Sombrero  Chino,  Rocas  Bainbridge,  SE  of 
Santiago,  0°21'S,  90°34'W,  31.iii.l970  (R.  Silberglied, 
MCZ);  Tower  Island,  Darwin's  Bay  [?],  0°17'N, 
89°59'W  (AMNH);  Tower  Islands,  0°20'N,  89°58'W, 
7.ivl925  (N.Y.  Zoological  Society,  AMNH);  W  coast 
of  Albemarle  Island,  0°11'N,  91°21'W,  9.iii.l935 
(AMNH). 

3.  Metepeira  grandiosa  grandiosa 
Chamberrm  &  Ivie 
Figures  22-28,  321;  Map  2 

Metepeira  grandiosa  Chamberlin  and  Ivie,  1941:  17, 
figs.  24—26,  9 .  Female  holotype  from  Ben  Lo- 
mond, California,  USA  in  the  AMNH,  examined. 

Metepeira  palomara  Chamberlin  and  Ivie,  1942:  72, 
figs.  200-204,  9,  6.  Female  holotype  and  para- 
types  from  Mt.  Palomar,  California,  in  the  AMNH, 
examined.  First  synonymized  by  Levi,  1977:  214. 

Metepeira  grandiosa  grandiosa: — Levi,  1977:  214, 
figs.  112-116,  ?,  (J. 

Note.  Levi  (1977)  opted  to  collapse  M.  paliistris, 
M.  grandiosa,  M.  alpina,  M.  dakota,  and  M.  palo- 
mara into  three  subspecies  of  M.  grandiosa,  with 
the  caveat  that  more  data  may  show  the  three  sub- 
species to  be  distinct  species.  D.  Buckle  (personal 
communication)  claims  that  his  own  recent  obser- 
vations suggest  that  M.  grandiosa  alpina  and  M. 
grandiosa  palustHs  are  separate  species.  However, 
since  the  bulk  of  M.  grandiosa  specimens  are  out- 
side of  the  geographic  range  of  this  revision,  and 
in  the  absence  of  molecular  data,  I  will  follow 
Levi's  (1977)  recommendation  and  leave  these  as 
separate  subspecies. 

Description.  Female  from  Los  Angeles, 
California,  USA.  Carapace  light  around 
eyes  with  inedian  white  line  extending  to 
thoracic  furrow  (Fig.  27).  Legs  ringed.  Fe- 
mur I  with  row  of  three  to  four  macrosetae 
on  anterior;  one  on  anteroventral.  Dorsum 
with  usual  folium,  though  lighter  and  more 
speckled  than  in  most  species  (Fig.  27). 
Venter  with  a  wide  longitudinal  white  line. 
Pair  of  white  spots  on  either  side  of  spi- 
racle (Fig.  28).  Sternum  entirely  black 
(Fig.  28).  Ratio  of  eye  diameters:  posterior 
medians  and  anterior  medians  1.0,  anterior 
laterals  1.2,  posterior  laterals  1.  Anterior 
median  eyes  separated  by  1.9  diameters, 
posterior  median  eyes  by  1.3,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  3  diameters  of  anterior  lateral  eyes,  lat- 
eral eyes  separated  by  0.4  their  diameters. 
Total  length  6.5  mm.   Carapace  3.2  mm 


24         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


long,  2.3  wide.  First  femur  3.5  mm,  patella 
and  tibia  3.7,  metatarsus  3.2,  tarsus  1.1. 
Second  patella  and  tibia  3.2  mm,  third  1.9, 
fourth  2.8. 

Male  from  Parque  Nacional  Sierra  San 
Pedro  Martir,  Baja  California  Norte,  Mex- 
ico. Carapace  as  in  female.  Legs  lightly 
ringed.  Macrosetae  on  femur  I  variable — 
usually  row  of  four  macrosetae  on  anterior 
side,  five  on  anteroventral  side.  Dorsum  as 
in  female  (Fig.  25).  Venter  and  sternum  as 
in  female  (Fig.  26).  Ratio  of  eye  diameters: 
posterior  medians  and  anterior  medians 
1.1,  anterior  laterals  1.5,  posterior  laterals 
1.5.  Anterior  median  eyes  separated  by  1.7 
diameters,  posterior  median  eyes  by  0.8, 
anterior  median  eyes  separated  from  an- 
terior laterals  by  2.1  diameters  of  anterior 
lateral  eyes,  lateral  eyes  separated  by  0.3 
their  diameters.  Total  length  3.6  mm.  Car- 
apace 1.9  mm  long,  1.5  wide.  First  femur 
2.9  mm,  patella  and  tibia  2.8,  metatarsus 
2.5,  tarsus  0.9.  Second  patella  and  tibia  2.3 
mm,  third  1.3,  fourth  1.9. 

Diagnosis.  The  epigynal  scape  of  M. 
grandiosa  grandiosa  is  shorter  and  stub- 
bier than  in  other  members  of  the  M.  foxi 
species  group  (compare  Fig.  24  with  Figs. 
9,  17,  31).  Unlike  M.  datona  and  M.  de- 
senderi  (Figs.  7,  15),  the  corkscrew  em- 
bolus is  tilted  up  with  a  heavy  ridge  (Fig. 
22),  but  unlike  M.  grandiosa  alpina  (Fig. 
29),  it  is  more  graceful  and  not  as  sharply 
bent.  The  posteroventral  edge  of  the  keel 
on  the  median  apophysis  in  M.  grandiosa 
grandiosa  is  not  as  pointed  (Fig.  22)  as  in 
M.  grandiosa  alpina  (Fig.  29)  but  not 
curved  as  in  M.  datona  and  M.  desenderi 
(Figs.  7,  15). 

Variation.  Body  length  of  females  varies 
from  5.4  to  8.5  mm;  males  from  3.5  to  5.1 
mm  (Levi,  1977). 

Natural  History.  Mature  specimens 
have  been  collected  from  March  to  Sep- 
tember (Fig.  321;  Levi,  1977)  in  yellow 
pine  forests  and  on  Eriogonum  fascictda- 
tum  bushes.  Elevations  range  from  sea  lev- 
el to  2,000  m. 

Distribution.    Coastal   mountainous   re- 


gions from  British  Columbia  to  Baja  Cal- 
ifornia Norte  (Levi,  1977,  map  2;  Map  2). 

Records  Examined.  MEXICO  Baja  California  Nor- 
te: Parque  Nacional  Sierra  San  Pedro  Martir, 
30°45'N,  115°13'W,  l.vii.l977  (C.  E.  Griswold,  CAS). 
USA  Arizona:  Sycamore  Canyon  [?]  Santa  Cruz  Co, 
31°28'N,  110°42'W,  9.ix.l978  (G.  F.  Knowlton, 
MCZ).  California:  Los  Angeles,  34°3'N,  118°15'W 
(Davidson,  MCZ);  Winchester,  Double  Butte, 
33°42'N,  117°5'W,  20.iv,1974  (W.  Icenogle,  MCZ). 

4.  Metepeira  grandiosa  alpina 
Chamberlin  and  Ivie 
Figures  29-35,  325;  Map  2 

Metepeira  dakota  Chamberlin  and  Ivie,  1942:  73,  figs. 
205—207,  ? ,  c? .  Male  holotype  and  female  paratype 
from  Noonan,  North  Dakota,  USA,  in  the  AMNH, 
examined.  Name  synonymized  by  Levi,  1977:  212— 
214. 

Metepeira  alpina  Chamberlin  and  Ivie,  1942:  74.  Fe- 
male holotype  and  female  paratypes  from  Fish 
Lake,  Utah,  USA,  in  tlie  AMNH,  examined. 

Metepeira  grandiosa  alpina: — Levi,  1977:  212-214, 
figs.  99,  100,  106-111,  9,6.  BrignoH,  1983:  276. 

ISlote.  As  first  revisor,  Levi  (1977)  chose  to  use 
the  name  M.  alpina  because  its  type  specimen  was 
collected  closer  to  the  center  of  the  subspecies  dis- 
tribution. 

Description.  Female  from  Charcas,  San 
Luis  Potosi,  Mexico.  Carapace  light 
around  eyes  with  lateral  posterior  exten- 
sions and  median  white  line  extending  td 
thoracic  furrow  (Fig.  34).  Legs  lightly 
ringed.  Femur  I  with  row  of  four  macro- 
setae on  anterior  side;  one  on  anteroven- 
tral side.  Dorsum  with  usual  folium, 
though  lighter  than  in  most  species  (Fig. 
34).  Venter  with  a  wide  longitudinal  white 
line.  Pair  of  white  spots  on  either  side  of 
spiracle  (Fig.  35).  Sternum  entirely  black 
(Fig.  35).  Ratio  of  eye  diameters:  posterior 
medians  and  anterior  medians  1.0,  anterior 
laterals  1.1,  posterior  laterals  1.1.  Anterior 
median  eyes  separated  by  1.7  diameters, 
posterior  median  eyes  by  0.7,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  2.8  diameters  of  anterior  lateral  eyes, 
lateral  eyes  separated  by  1.1  their  diame- 
ters. Total  length  6.7  mm.  Carapace  3  mm 
long,  2.4  wide.  First  femur  3.5  mm,  patella 
and  tibia  3.8,  metatarsus  3.4,  tarsus   1.2. 


Metepeira  •  Piel        25 


desenderi 

(2) 


grandiosa  grandiosa 
(3) 


m^  grandiosa  alpina 

V  (4) 


Figures  15-21.  Metepeira  desenderi  Baert  (sp.  2;  0°38'S,  90°23'W).  15,  male  palpus,  mesal.  16,  epigynum,  posterior.  17, 

epigynum,  ventral.  18,  male,  dorsal.  19,  male,  ventral.  20,  female,  dorsal.  21,  female,  ventral. 

Figures  22-28.  Metepeira  grandiosa  grandiosa  Chamberlin  and  Ivie  (sp.  3  [22,  25,  26]  30°45'N,  115°13'W;  [23,  24,  27,  28] 

34°3'N,  118°15'W).  22,  male  palpus,  mesal.  23,  epigynum,  posterior.  24,  epigynum,  ventral.  25,  male,  dorsal.  26,  male,  ventral. 

27,  female,  dorsal.  28,  female,  ventral. 

Figures  29-35.  Metepeira  grandiosa  alpina  Chamberlin  and  Ivie  (sp.  4  [29,32,33]  25°56'N,  105°22'W;  [30,31,34,35]  23°8'N, 

101°7'W).  29,  male  palpus,  mesal.  30,  epigynum,  posterior.  31,  epigynum,  ventral.  32,  male,  dorsal.  33,  male,  ventral.  34,  female, 

dorsal.  35,  female,  ventral. 

Scale  bars:  dorsum  and  venter  figures  1 .0  mm. 


26         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


10000 


1000 


■I    100  ; 


°  vigilax 
X  rectangula 
•  cajabamaba 


JQ  I    I    I    I  ''I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I    I 

-45         -35         -25         -15  -5  5  15  25 

Latitude  (Degrees  North) 

Figure  36.  The  elevation  of  collection  localities  for  M.  vigilax, 
M.  rectangula,  and  M.  cajabamba  at  their  corresponding  lati- 
tudes. Species-specific  altitudes  appear  to  decrease  with  dis- 
tance from  the  equator.  Elevations  estimated  from  NOAA  da- 
tabase of  5-  by  5-minute  geographic  tiles.  Regression  line  of 
M.  vigilax  does  not  include  data  points  north  of  the  equator. 
Symbols:  M.  vigilax  (o),  M.  rectangula  [X],  M.  cajabamba  [•]. 


Second  patella  and  tibia  3.3  mm,  third  1.9, 
fourth  2.9. 

Male  from  Santa  Maria  del  Oro,  Duran- 
go,  Mexico.  Carapace  yellowish  brown, 
light  around  eyes.  Median  white  triangular 
mark  anterior  to  thoracic  furrow  (Fig.  32). 
Legs  same  color  as  carapace.  Macrosetae 
on  femur  I  variable — usually  row  of  four 
macrosetae  on  anterior  side,  five  to  six  on 
anteroventral  side.  Dorsum,  venter,  and 
sternum  as  in  female  (Figs.  32,  33).  Ratio 
of  eye  diameters:  posterior  medians  and 
anterior  medians  1.0,  anterior  laterals  1.4, 
posterior  laterals  1.2.  Anterior  median 
eyes  separated  by  1.5  diameters,  posterior 
median  eyes  by  0.8,  anterior  median  eyes 
separated  from  anterior  laterals  by  2.3  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.3  their  diameters.  To- 
tal length  5.2  mm.  Carapace  2.5  mm  long, 
2  wide.  First  femur  4.5  mm,  patella  and 
tibia  4.6,  metatarsus  4.5,  tarsus  1.3.  Sec- 
ond patella  and  tibia  3.6  mm,  third  1.8, 
fourth  2.8. 

Diagnosis.  The  epigynal  scape  of  M. 
grandiosa  alpina  has  the  widest  base  of  all 
members  of  the  M.  foxi  species  group,  and 
the  lobes  surrounding  the  central  depres- 
sion are  soiuewhat  fatter  (compare  Fig.  31 
witli  Figs.  9,  17,  24,).  Unlike  M.  datona 
and  M.  desenderi  (Figs.  7,  15),  the  cork- 


screw embolus  is  tilted  up  with  a  heavy 
ridge  (Fig.  29),  but  unlike  M.  grondiosa 
grandiosa  (Fig.  22),  it  is  thicker  and  with 
a  sharper  L-shape  bend  instead  of  a  more 
cui'ved  L-shape.  The  posteroventral  edge 
of  the  keel  on  the  median  apophysis  in  M. 
grandiosa  alpina  is  pointier  (Fig.  29)  than 
in  M.  grandiosa  grandiosa  (Fig.  22)  and 
not  curved  as  in  M.  datona  and  M.  desen- 
deri (Figs.  7,  15). 

Variation.  Body  length  of  females  varies 
from  4.0  to  6.8  mm;  males  from  3.1  to  5.3 
mm  (Levi,  1977). 

Natural  History.  According  to  Levi 
(1977),  mature  specimens  have  been  col- 
lected from  June  to  August  in  meadows  of 
bunchgrass,  browsed  aspen,  oak,  juniper, 
and  sagebrush  (Fig.  325).  Elevations  are  at 
around  2,000  m. 

Distribution.  North  American  Rockies 
from  southern  Alberta  and  Saskatchewan 
to  Central  Mexico  (Levi,  1977,  map  2; 
Map  2). 

Records  Examined.  MEXICO  Durango:  Santa  Ma- 
ria del  Oro,  25°56'N,  105°22'W,  28.vii.1947  (W.  J. 
Gertsch,  AMNH).  San  Luis  Potosi:  Charcas,  moun- 
tain side,  23°8'N,  101°7'W,  7.vii.l934  (MCZ).  USA. 
Colorado:  Cimarron,  38°27'N,  107°33'W,  21.vii.l959 
(H.  W.  &  L.  Levi,  MCZ);  Hayden  Creek,  Sangre  de 
Cristo  Mtns.,  38°25'N,  105°35'W  [?],  Il.vii.l961  (H. 
W.  &  L.  Levi,  MCZ).  Utah:  SE  shore  of  Bear  Lakei 
41°59'N,  111°20'W,  3.vii.l978  (G.  F.  Knovi^lton, 
MCZ). 

Metepeira  vigilax  Group 

Spiders  in  the  M.  vigilax  group  (Mete- 
peira cajabamba,  Metepeira  glome rabilis, 
Metepeira  vigilax,  Metepeira  rectangula) 
are  characterized  by  large,  sclerotized  epi- 
gynal openings,  not  unlike  a  snails  shell 
(Figs.  40,  48,  55,  62),  and  long,  robust, 
emboli  with  large  scooplike  basal  embolic 
apophyses  (Figs.  38,  46,  53,  60). 

5.  Metepeira  cajabamba  new  species 
Figures  36,  38-45,  315;  Map  3 

Holotype.  Male  from  Cajabamba,  Cajamarca,  Peru, 
25. Lx.  1955,  W  Weyi-auch,  in  CAS.  The  specific 
name  is  a  noun  in  apposition  after  the  localit)'. 

Description.  Female  paratype  from  Ca- 
jabamba, Cajamarca,  Pei-u.  Carapace  dark 


Metepeira  •  Piel        27 


brown,  light  around  eyes  with  lateral  pos- 
terior extensions  (Fig.  43).  Proximal  two- 
thirds  of  femora  white,  remainder  dark 
browii.  Remaining  articles  lightly  annulat- 
ed.  Femur  I  with  row  of  three  macrosetae 
on  anterior  side;  none  on  anteroventral 
side.  Dorsal  folium  dark;  white  fleur-de-lis 
pattern  reduced  with  thin  branches  (Fig. 
43).  Venter  of  abdomen  black  with  wide 
white  median  line.  Pair  of  small  white 
spots  on  either  side  of  colulus  (Fig.  44). 
Sometimes  the  posterior  end  of  median 
line  ends  in  anchor  shape  (Fig.  45).  Ster- 
num black,  sometimes  with  (Fig.  44)  or 
without  (Fig.  45)  median  white  line.  Ratio 
of  eye  diameters:  posterior  medians  and 
anterior  medians  1.1,  anterior  laterals  1.4, 
posterior  laterals  1.2.  Anterior  median 
eyes  separated  by  1.6  diameters,  posterior 
median  eyes  by  0.9,  anterior  median  eyes 
separated  from  anterior  laterals  by  3  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.2  their  diameters.  To- 
tal length  6.4  mm.  Carapace  3  mm  long, 
2.3  wide.  First  femur  3.3  mm,  patella  and 
tibia  3.5,  metatarsus  2.9,  tarsus  0.9.  Sec- 
ond patella  and  tibia  2.9  mm,  third  1.7, 
fourth  2.7. 

Male  holotype.  Carapace  dark  brown, 
white  around  eyes  with  lateral  posterior 
extensions  and  median  white  mark  (Fig. 
41).  Proximal  third  of  femora  white,  re- 
mainder dark  brown.  Remaining  articles 
lightly  annulated.  Femur  I  with  row  of 
three  macrosetae  on  anterior  side;  one  on 
anteroventral  side.  Dorsal  folium  dark; 
white  fleur-de-lis  pattern  indistinct  with 
thin  and  broken  branches  (Fig.  41).  Venter 
of  abdomen  black  with  median  white 
mark.  Pair  of  small  white  spots  on  either 
side  of  spiracle  and  colulus  (Fig.  42).  Ster- 
num black,  often  with  median  white  line 
(Fig.  42).  Ratio  of  eye  diameters:  posterior 
medians  and  anterior  inedians  1.0,  anterior 
laterals  1.3,  posterior  laterals  1.  Anterior 
median  eyes  separated  by  1.5  diameters, 
posterior  median  eyes  by  0.9,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  2.3  diameters  of  anterior  lateral  eyes, 
lateral  eyes  separated  by  0.3  their  diame- 


ters. Total  length  4  mm.  Carapace  1.9  mm 
long,  1.5  wide.  First  femur  3  mm,  patella 
and  tibia  3,  metatarsus  2.7,  tarsus  0.9.  Sec- 
ond patella  and  tibia  2.4  mm,  third  1.2, 
fourth  1.8. 

Diagnosis.  Female  M.  cajahamba  and 
M.  glomerabilis  differ  from  the  other  spe- 
cies in  the  M.  vigilax  group  (M.  vigilax  and 
M.  rectangula)  by  the  smaller  and  more 
tubelike  openings  to  the  epigynum  (com- 
pare Figs.  40,  48  with  Figs.  55,  62).  Fe- 
male M.  cajabamha  differs  froin  M.  glom- 
erabilis by  having  epigynal  openings  that 
are  more  oval  (Fig.  40)  than  round  (Fig. 
48)  when  viewed  ventrally,  having  the  epi- 
gynal openings  farther  apart  (compare  Fig. 
40  with  Fig.  48),  and  by  having  the  scler- 
otized  tubelike  openings  more  anteriorly 
directed  (Fig.  40)  than  parallel  to  the  epi- 
gynal groove  (Fig.  48).  Male  M.  cajabarnba 
and  M.  glomerabilis  differ  from  other  spe- 
cies in  the  M.  vigilax  group  by  the  smaller, 
thinner,  and  more  graceful  emboli  (com- 
pare Figs.  38,  46  with  Figs.  53,  60).  Me- 
tepeira cajahamba  differs  from  M.  glom- 
erabilis by  lacking  a  keel  on  the  median 
apophysis,  an  only  slightly  fatter  embolus, 
and  having  a  normal  embolus  cap,  in  con- 
trast to  a  winged  embolus  cap  (Fig.  46).  A 
larger  portion  of  the  prosoma  is  dark  in  M. 
cajahamba  (Fig.  43)  as  compared  to  M. 
glom,erabilis  (Fig.  51). 

Variation.  Average  body  length  of  nine 
females  examined  6.2  mm,  range  5.4  to  7.5 
mm.  Average  body  length  of  four  males 
examined  4  mm,  range  3.7  to  4.9  mm. 

Natural  History.  With  the  exception  of 
two  specimens,  mature  M.  cajabarnba 
specimens  have  been  collected  in  May 
through  October  (Fig.  315).  Altitudes  of 
collection  localities  appear  to  correlate 
steeply  with  latitude  (Fig.  36).  Median  el- 
evation, about  500  m,  with  a  range  from 
near  sea  level  to  3,500  m. 

Distribution.  Ecuador  and  Peru  (Map 
3). 

Records  Examined.  PERU  Ancash:  Callejon  de 
Huaylas,  9°10'S,  77°45'W,  15.viii.l988  (V.  and  B. 
Roth,  CAS).  Cajamnrca:  Cajabarnba,  12.427,  7°37'S, 
78°3'W,  25.ix.1955  (W.  Weyrauch,  CAS).  La  Libet-tad: 


28         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


Cerro  Campana,  La  Cumbre,  8°1'S,  79°5'W, 
10.x.  1966  (A.  F.  Archer,  AMNH).  Liiiia:  20  km  E  An- 
con,  11°47'S,  76°59'W,  l.xi.l953  (W.  Weyrauch, 
CAS);  23  mi  N  Pativilca,  10°42'S,  77°47'W,  15.1.1955 
(E.  I.  Schlinger  and  E.  S.  Ross,  CAS);  5  km  NW 
Chilca,  12°29'S,  76°48'W,  12.ix.l954  (E.  I.  Schlinger 
and  E.  S.  Ross,  CAS);  between  moudis  of  Rio  Chillon 
and  Ancon  [?],  11°54'S,  77°7'W,  5.viii.l953  (M. 
Koepcke  and  Koepcke,  MUSM);  Cajacay  [?],  Rio 
Fortaleza,  10°40'S,  77°52'W,  6.iii.l956  (W.  Weyrauch, 
CAS);  Canta,  Rio  Chillon,  11°28'12"S,  76°37'23"W, 
12.V.1951  (W.  Weyrauch,  CAS);  Cerro  Caracoles, 
12°23'S,  76°45'W  15.b£.1951  (W.  Weyi-auch,  CAS); 
Lima,  12°3'S,  77°3'W  (K.  Jelski  and  Stolzman,  PAN); 
Lomas  de  Iguanil  [?]  (Huaral),  11°29'51"S, 
77°12'12"W,  14.vi.l986  (D.  Silva,  MUSM). 

6.  Metepeira  glomerabilis  (Keyserling) 
Figures  37,  46-52,  314;  Map  3 

Epeira  glomerabilis  Keyserling,  1892:  154,  fig.  113, 
9 ,  <S .  Male  and  female  syntypes  from  Taquara,  Rio 
Grande  do  Sul  and  Serra  Vermelha,  Rio  de  Janeiro, 
Brazil,  in  BMNH,  examined.  Male  here  designated 
lectotype. 

Araneus  glomerabilis: — Petrunkevitch,  1911:  294. 
Roewer,  1942:  843. 

Aranea  .santa  Chamberlin,  1916:  254,  pi.  19,  fig.  10, 
9 .  Female  holotype  from  Santa  Ana,  3,000  m,  Cuz- 
co,  Peru,  in  MCZ,  examined.  NEW  SYNONYMY. 

Metepeira  santa: — Chamberlin  and  Ivie,  1942:  67. 
Platnick,  1993:  449. 

Metepeira  glomerabilis: — Chamberlin  and  Ivie,  1942: 
68,  fig.  181. 

Metazygia  gloiTierabilis: — Levi,  1991:  179.  Platnick, 
1993:  448.  Erroneous  transfer. 

Description.  Female  from  Punapi,  Tun- 
gurahua,  Ecuador.  Carapace  brown;  white 
arrowhead  mark  between  eye  region  and 
thoracic  furrow  (Fig.  51).  Legs  dusky 
brown,  hghter  on  ventral  surfaces  of  fem- 
ora I  and  II;  lighter  on  dorsal  surfaces  of 
patellae,  tibiae,  and  metatarsi  I  and  II.  Fe- 
mur I  with  row  of  two  to  three  macrosetae 
on  anterior  side;  none  on  anteroventral 
side.  Dorsal  folium  dark,  speckled  white, 
and  white  fleur-de-lis  pattern  with  thin 
branches  (Fig.  51).  Venter  black  with 
short,  wide,  white  median  line  ending  in  a 
T-shape;  pair  of  small  white  spots  on  either 
side  of  spiracle  (Fig.  52).  Sternum  black 
with  median  white  line  widening  anteriorly 
(Fig.  52).  Ratio  of  eye  diameters:  posterior 
medians  and  anterior  medians  1.0,  anterior 
laterals  1.3,  posterior  laterals  1.2.  Anterior 
median  eyes  separated  by  1.3  diameters, 


posterior  median  eyes  by  0.9,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  1.7  diameters  of  anterior  lateral  eyes, 
lateral  eyes  separated  by  0.2  their  diame- 
ters. Total  length  5  mm.  Carapace  2.4  mm 
long,  1.7  wide.  First  femur  2.2  mm,  patella 
and  tibia  2.3,  metatarsus  1.9,  tarsus  0.9. 
Second  patella  and  tibia  1.9  mm,  third  1.2, 
fourth  1.8. 

Male  from  Puiiapi,  Tungurahua,  Ecua- 
dor. Carapace  as  in  female,  except  for  a 
greater  separation  between  white  colora- 
tion around  eyes  and  arrowhead  mark 
(Fig.  49).  Legs  light  tan,  dark  distally  on 
femora.  Femur  I  with  row  of  three  macro- 
setae  on  anterior  side;  four  on  anteroven- 
tral side.  Dorsal  folium  and  venter  as  in 
female  (Figs.  49,  50).  Sternum  black  with 
partly  broken  median  white  line  widening 
anteriorly  (Fig.  50).  Ratio  of  eye  diame- 
ters: posterior  medians  and  anterior  me- 
dians 1.1,  anterior  laterals  1.3,  posterior 
laterals  1.2.  Anterior  median  eyes  separat- 
ed by  1.8  diameters,  posterior  median  eyes 
by  1.2,  anterior  median  eyes  separated 
from  anterior  laterals  by  2  diameters  of  an- 
terior lateral  eyes,  lateral  eyes  separated  by 
0.3  their  diameters.  Total  length  3.5  mm. 
Carapace  1.7  mm  long,  1.4  wide.  First  fe- 
mur 2.5  mm,  patella  and  tibia  2.4,  meta- 
tarsus 2.3,  tarsus  0.9.  Second  patella  and 
tibia  1.9  mm,  third  1.0,  fourth  1.7. 

Diagnosis.  Female  M.  ^omerabilis  and 
M.  cajabamba  differ  from  the  other  spe- 
cies in  the  M.  vigilax  group  (M.  vigilax  and 
M.  rectangula)  by  the  smaller  and  more 
tubelike  openings  to  the  epigynum  (com- 
pare Figs.  40,  48  with  Figs.  55,  62).  Me- 
tepeira glomerabilis  differs  from  M.  caja- 
bamba by  having  epigynal  openings  that 
are  more  round  (Fig.  48)  than  oval  (Fig. 
40)  when  viewed  ventrally,  having  the  epi- 
gynal openings  closer  together  (compare 
Fig.  48  with  Fig.  40),  and  by  having  the 
sclerotized  tubelike  openings  more  parallel 
to  the  epigynal  groove  (Fig.  48)  than  an- 
teriorly directed  (Fig.  40).  Male  M.  glom- 
erabilis and  M.  cajabamba  differ  from  oth- 
er species  in  the  M.  vigilax  group  by  the 
smaller,  thinner,  and  more  graceful  emboli 


Metepeiba  •  Piel 


29 


(compare  Figs.  38,  46  with  Figs.  53,  60). 
Metepeira  glomerabilis  differs  from  M.  ca- 
jabaniba  by  having  a  keel  on  the  median 
apophysis,  an  only  slightly  slimmer  embo- 
lus, and  a  larger,  winged  embolus  cap 
(compare  Fig.  46  with  Fig.  38).  A  larger 
portion  of  the  prosoma  is  white  in  M. 
glo77ierabilis  (Fig.  51),  compared  to  M.  ca- 
jabaniba  (Fig.  43).  In  addition,  the  mar- 
gins of  the  folium,  particularly  in  the  male, 
are  whiter  than  in  other  species  in  the 
group  (compare  Fig.  49  and  Keyserling 
[1892,  fig.  113b]  with  Figs.  41,  56,  63). 

Variation.  Average  body  length  of  28  fe- 
males examined  5.1  mm,  range  3.8  to  7.3 
mm.  Average  body  length  of  10  males  ex- 
amined 3.1  mm,  range  2.4  to  4.3  mm.  The 
base  of  the  embolus  varies  from  relatively 
thin  (Fig.  46)  to  somewhat  thicker,  as  in 
M.  cajabamba  (Fig.  38). 

Natural  History.  At  first  it  would  appear 
that  this  species  is  not  seasonal — mature 
specimens  have  been  collected  throughout 
the  year  (Fig.  315).  However  on  closer  in- 
spection, there  seems  to  be  a  seasonal  shift 
with  elevation:  mature  spiders  are  found 
at  low  altitudes  (0—500  m)  between  March 
and  October,  at  medium  altitudes  (500— 
1,500  m)  between  August  and  March,  and 
at  high  altitudes  (1,500-4,000  m)  between 
December  and  June  (Fig.  37).  In  coastal 
regions,  R.  Baptista  (personal  communi- 
cation) reports  that  this  species  forms 
small  aggregations  of  two  to  10  spiders. 

Distribution.  Colombia,  Ecuador,  Peru, 
Bolivia,  Paraguay,  and  southern  Brazil 
(Map  3).  Elevations  range  froin  sea  level 
to  4,000  m. 

Records  Examined.  BOLIVIA  Chuquisaca:  Mon- 
teagudo,  19°49'S,  63°59'W,  24.xii.1984  (L.  E.  Pena, 
AMNH).  BRAZIL  EspiHto  Santo:  Fazenda  Santa 
Maria  [?],  Apiaca,  21°4'S,  41°25'W,  22. ix.  1985  (R.  L. 
C.  Baptista,  MZSP).  Mato  Grosso:  Chavantina, 
14°40'S,  52°21'W,  15.vi.l947  (J.  C.  Carvalho,  MNRJ); 
Fazenda  Cei-vo,  Tres  Lagoas,  20°48'S,  51°43'W, 
18. ix.  1964  (Exp.  Depto.  ZooL,  MZSP);  Utiariti, 
13°2'S,  58°17'W,  15.vlii.l961  (Lenks,  MZSP).  Minas 
Gerais:  Lavras,  21°14'S,  45°0'W,  20.x.  1978  (W.  Don 
Fronk,  MCZ);  Pedra  Azul,  16°1'S,  41°16'W, 
15.xii.l970  (F.  M.  Oliveira,  AMNH);  Peti  Forest  Res. 
[?]  Santa  Barbara,  on  bushes  in  cerrado,  19°56'S, 
43°24'W,  28.viii.1986  (R.  L.  C.  Baptista,  MZSP).  Pa- 


Peru,  Ecuador,  Colombia 
Brazil,  Paraguay,  Bolivia 


50      100     150    200     250     300    350     400    450    500     550 
Collection  Day  (since  Jan  1) 

Figure  37.  The  elevation  of  collection  localities  for  mature  spi- 
ders of  M.  glomerabilis  on  specific  days  of  the  year  between 
1939  and  1990.  There  appears  to  be  a  shift  in  seasonal  mat- 
uration times  that  corresponds  better  with  elevation  than  with 
latitude.  Elevations  estimated  from  NOAA  database  of  5-  by 
5-minute  geographic  tiles. 

Scale  of  abscissa:  1 00  =  April  9;  300  =  October  27;  500  = 
May  15  of  the  following  year. 

Symbols:  Peru,  Ecuador,  Colombia  [■];  Brazil,  Paraguay,  Bo- 
liva  [♦]. 


rand:  Praia  do  Leste,  Paranagua,  2.5°46'S,  48°31'W, 
4.V.1967  (P.  Biasi,  MZSP).  Rio  de  Janeiro:  Guaratiba, 
22°58'S,  42°48'W,  28.viii.1976  (J.  A.  R  Dutra,  MZSP); 
Ilha  de  Santana,  Macae,  22°25'S,  41°44'W,  18.x.  1986 
(R.  L.  C.  Baptista,  MZSP);  Itaipu,  Niteroi,  22°56'S, 
43°5'W,  20.ivl985  (R.  L.  C.  Baptista,  MZSP).  Rio 
Grande  do  Norte:  Fazenda  Canaan  [?],  Macaiba, 
5°51'S,  35°21'W,  15.ix.l951  (M.  Alvorenga,  MZSP). 
Rondonia:  Vila  Rondonia,  10°52'S,  61°57'W,  9.ii.l961 
(Pereira  &  Machado,  MZSP).  Santa  Catarina:  Nova 
Teutonia,  27°3'S,  52°24'W,  12.vl949  (SMF).  Sao 
Paulo:  Barueri,  23°31'S,  46°53'W,  13.iii.l966  (K.  Len- 
ko,  MZSP);  Campos  do  Jordao,  22°44'S,  45°35'W, 
15.iii.l945  (Wygod,  MZSP);  Institute  Oceanografico, 
Ubatuba,  2.3°26'S,  45°4'W,  1.5.vl967  (P  Montouchat, 
MZSP);  km  1  Rod,  Rio  Santos  [?],  Ubatuba,  23°26'S, 
45°4'W,  12.X.1985  (R.  L.  C.  Baptista,  MZSP), 
13.X.1985  (R.  L.  C.  Baptista,  MZSP);  km  3  Rod,  Rio 
Santos  [?],  Ubatuba,  23°26'S,  45°4'W,  6.i.l985  (R.  L. 
C.  Baptista,  MZSP).  COLOMBIA  Cundinamarca: 
Sabana  de  Bogota,  4°43'N,  74°10'W,  10.xii.l990  (C. 
Valderrama,  CV).  ECUADOR  Guaijas:  Guayaquil, 
2°10'S,  79°.54'W,  18.iii.l942  (H.  E.  F.  &  D.  E.  F, 
CAS),  22.iii.1942  (Landis,  CAS).  Pichlncha:  1.5  mi  N 
Quito,  0°0'N,  78°30'W,  23.ii.1955  (E.  I.  Sehlinger  & 
E.  S.  Ross,  CAS).  Tungurahua:  Baiios,  1°24'S, 
78°25'W,  15.ivI939  (W.  C.  Macintyre,  MCZ);  Puiiapi, 
1°22'S,  78°28'W,  19.vi.l943  (D.  L.  F.  &  H.  E.  F, 
CAS).  PARAGUAY  Alto  Parana:  Taguararaya  [?], 
25°30'S,  .54°50'W  (AMNH).  Caazapa:  Villa  Pastoreo, 
25°53'S,  55°45'W  (D.  Wees,  MCZ).  PERU  Cajamar- 
ca:  Cajamarca,  7°10'S,  78°31'W,  15.ii.l942  (W.  Wey- 
rauch,  AMNH).  Lanihaijeque:  10  km  S  Chiclayo, 
7°59'S,  77°17'W,  19.iii.l951  (E.  S.  Ross  &  Michel- 
bacher,  CAS).  Piura:  Cerro  Prieto,  La  Brea,  4°41'S, 
81°6'W  (CAS);  Higueron  (Las  Lomas)  [?],  4°19'S, 
80°26'W,  29.vii.194I  (D.  L.  F  &  H.  E.  F,  CAS). 
VENEZUELA  Sucre:  1  km  S  Villa  Frontado,  Rd.  to 


30         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


Caripe,  10°27'N,  63°37'W,  12.ii.l984  (J.  Coddington, 

USNM). 

7.  Metepeira  vigilax  (Keyserling) 
Figures  36,  53-59,  327;  Map  4 

Epeira   vigilax   Keyserling,    1893:   211,   fig.    156,    6. 

Male  holotype  from  Taquara,  Rio  Grande  do  Sul, 

Brazil,  in  BMNH,  examined. 
Araneus  vigilax: — Petrunkeviteh,  1911:  324.  Roewer, 

1942:  856. 
Metepeira  dominicana  Archer,  1965:  132,  figs.  12,  18, 

$  .  Female  holotype  from  west  of  Bani,  Dominican 

Republic,  in  AMNH.  Holotype  lost.  Brignoli,  1983: 

275.  NEW  SYNONYMY. 
Metepeira  vigilax: — Levi,  1991:  180.  Platnick,  1993: 

449. 

Note.   Although  the  type  for  M.   dominicana  is 

lost,  the  name  has  been  identified  by  using  Archer's 

(1965)  description  and  illustration. 

Description.  Female  from  Trujillo,  west 
of  Bani,  Dominican  Republic.  Carapace 
light  around  eyes  with  lateral  posterior  ex- 
tensions (Fig.  58).  Legs  dark,  light  rings  on 
proximal  ends  of  leg  articles.  Femur  I  with 
ro\v  of  four  macrosetae  on  anterior  side; 
three  on  anteroventral  side.  Dorsal  folium 
darker  than  in  most  species;  fleur-de-lis 
usually  reduced  to  two  white  spots  (Fig. 
58).  Venter  brownish  gray  with  lighter 
margins.  Wide,  short,  median  white  line 
with  pair  of  white  spots  on  either  side  of 
spiracle  (Fig.  59).  Sternum  brownish  black 
with  wide,  white  line  widening  anteriorly 
(Fig.  59).  Ratio  of  eye  diameters:  posterior 
inedians  and  anterior  medians  1.2,  anterior 
laterals  1.2,  posterior  laterals  1.1.  Anterior 
median  eyes  separated  by  1.8  diameters, 
posterior  inedian  eyes  by  1.1,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  3.5  diameters  of  anterior  lateral  eyes, 
lateral  eyes  separated  by  0.1  their  diame- 
ters. Total  length  9.2  mm.  Carapace  3.9 
mm  long,  3.2  wide.  First  femur  4.1  mm, 
patella  and  tibia  4,  metatarsus  3.3,  tarsus 

1.2.  Second  patella  and  tibia  3.4  mm,  third 

2.3,  fourth  3.4. 

Male  froin  same  locality  as  female. 
Black  carapace  with  white  around  eyes  and 
extending  posteriorly;  white  wedge  mark 
in  center  (Fig.  56).  Legs  ringed  like  fe- 
male. Femur  I  with  row  of  three  luacro- 
setae  on  anterior  side;  three  on  anterov- 


entral side.  Dorsum  and  venter  as  in  fe- 
male (Figs.  56,  57).  Median  white  line  may 
be  limited  to  posterior  end  of  sternum 
(Fig.  57).  Ratio  of  eye  diameters:  posterior 
medians  and  anterior  inedians  0.9,  anterior 
laterals  1.1,  posterior  laterals  1.  Anterior 
median  eyes  separated  by  1.6  diameters, 
posterior  median  eyes  by  1.3,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  2  diameters  of  anterior  lateral  eyes,  lat- 
eral eyes  separated  by  0.2  their  diameters. 
Total  length  3.9  mm.  Carapace  2  mm  long, 
1.5  wide.  First  femur  2.6  mm,  patella  and 
tibia  2.5,  metatarsus  2.4,  tarsus  0.9.  Sec- 
ond patella  and  tibia  2.1  mm,  third  1.2, 
fourth  1.7. 

Diagnosis.  Female  M.  vigilax  differ 
from  those  of  other  species  in  the  M.  vi- 
gilax species  group  by  the  shape  of  the 
epigynal  openings:  from  a  ventral  view  the 
openings  are  oval  and  angled  inward  pos- 
teriorly (Fig.  55);  from  a  posterior  view, 
the  edges  of  the  openings  are  more  par- 
allel to  the  body  (Fig.  61)  as  opposed  to 
more  pei^pendicular  to  the  body  (Figs.  39, 
61).  Male  M.  vigilax  differ  from  other  spe- 
cies because  the  embolus  is  larger  and 
more  robust  (compare  Fig.  53  with  Figs. 
38,  46,  60);  the  two  flagella  on  the  median 
apophysis  are  of  more  similar  width  (com- 
pare Fig.  53  with  Fig.  38),  and  the  keel  on 
the  median  apophysis  is  slim  and  feather- 
shaped  (Fig.  38),  in  contrast  to  arrow- 
shaped  (Figs.  46,  60)  or  absent  (Fig.  38). 
The  dorsal  folium  differs  from  other  Me- 
tepeira species  by  having  a  wide  black  me- 
dian stripe  at  the  posterior  end  of  the  ab- 
domen (Fig.  58).  In  Brazilian  and  Bolivian 
specimens  this  stripe  often  extends  all  the 
way  to  the  black  anterior  shoulders  of  the 
dorsum,  forming  a  wide  T-shape  mark. 

Variation.  Specimens  from  Argentina 
tend  to  be  more  lightly  pigmented  than 
those  from  more  northern  localities.  White 
markings  on  the  eye  region  of  Brazilian 
and  Bolivian  specimens  surround  only  the 
lateral  eyes,  in  contrast  to  those  on  His- 
paniolan  specimens,  which  cover  the  en- 
tire eye  region. 

Natural    History.    Mature    adults    have 


Metepeira  •  Piel        31 


cajabamba 

(5) 


Figures  38^5.  Metepeira  cajabamba  new  species  (sp.  5  [38-44]  7°37'S,  78°3'W;  [45]  9°10'S,  77°45'W).  38,  male  palpus, 

mesal.  39,  epigynum,  posterior.  40,  epigynum,  ventral.  41 ,  male,  dorsal.  42,  male,  ventral.  43,  female,  dorsal.  44,  female,  ventral. 

45,  female,  ventral. 

Figures  46-52.  Metepeira  glomerabilis  (Keyserling)  (sp.  6;  1°22'S,  78°28'W).  46,  male  palpus,  mesal.  47,  epigynum,  posterior. 

48,  epigynum,  ventral.  49,  male,  dorsal.  50,  male,  ventral.  51,  female,  dorsal.  52,  female,  ventral. 

Figures  53-59.  Metepeira  vigilax  (Keyserling)  (sp.  7  [53-57]  18°27'N,  72°17'W;  [58,59]  18°17'N,  70°22'W).  53,  male  palpus, 

mesal.  54,  epigynum,  posterior.  55,  epigynum,  ventral.  56,  male,  dorsal.  57,  male,  ventral.  58,  female,  dorsal.  59,  female,  ventral. 

Scale  bars:  dorsum  and  venter  figures  1 .0  mm. 


32         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


been  collected  throughout  the  year  except 
Januaiy,  February,  and  March  (Fig.  327). 
Spiders  have  been  found  among  electric 
wires  four  meters  above  ground.  Locality 
elevations  range  from  near  sea  level  to 
1,400  m  and  follow  an  ecological  zone  that 
decreases  in  elevation  with  distance  from 
the  equator  (Fig.  36).  Over  equivalent  lat- 
itudes, M.  vigilax  lives  at  less  than  one- 
tenth  of  the  elevation  of  M.  rectangula 
(Fig.  36). 

Distribution.  Hispaniola,  Bolivia,  Brazil, 
and  coastal  Argentina  (Map  4).  The  dis- 
junct distribution  between  Hispaniolan 
and  South  American  populations  may  be 
due  to  human-assisted  migration. 

Records  Examined.  ARGENTINA  Buenos  Aires: 
Celsya-Pereyra  [?],  34°50'S,  58°6'W  (MACN);  Ze- 
laya,  34°21'S,  58°52'W  (MACN).  BOLIVIA  La  Paz: 
Apolo,  14°43'S,  68°31'W,  10.viii.l989  (L.  E.  Pena, 
AMNH).  Santa  Cruz:  Estacion  Robore,  above  creek, 
18°20'S,  59°45'W,  27.ix.1955  (Azambuya,  CAS). 
BRAZIL  Espirito  Santo:  Fazenda  Santa  Maria  [?], 
Apiaca,  21°4'S,  41°25'W,  14.V.1988  (R.  L.  C.  Baptista, 
MZSP).  Rio  Grande  do  Sul:  Sao  Leopoldo,  29°46'S, 
5r9'W,  14.vi.l964  (Celia  Valle,  MZSP).  DOMINI- 
CAN REPUBLIC  Aziia:  El  Pueriio,  Majagual  and 
Peralta,  I8°34'N,  70°47'W,  10.xi.l979  (E.  Marcano, 
MNSD).  Prov.  Tnijillo  Vdldez:  W  Bani,  18°17'N, 
70°22'W,  8.viii.l958  (A.  F.  Archer  &  E.  de  Boyrie 
Moya,  AMNH).  HAITI  Departement  de  L' Quest: 
Kenscoff,  18°27'N,  72°17'W,  15.xii.l929  (J.  C.  Myers, 
AMNH),  17.iv.l935  (AMNH). 

8.  Metepeira  rectangula  (Nicolet) 
Figures  36,  60-66,  306;  Map  4 

Epeira  rectangula  Nicolet,  1849:  500,  female  holo- 

type  from  Valdivia,  Chile,  in  MNHN. 
Metepeira    labyrinthea: — Petrunkevitch,    1911:    298. 

Roewer,  1942:  868.  Bonnet,  1957:  2821.  Erroneous 

synonymy. 
Metepeira  rectangulata: — Chamberlin  and  Ivie,  1942: 

71.  Unjustified  emendation. 

Note.  The  name  was  identified  using  drawings  of 

the  holotype  (H.  W.  Levi,  personal  illustrations). 

Description.  Female  from  Angol,  Mal- 
leco,  Chile.  Carapace  reddish  brown  with 
long  white  setae  behind  lateral  eyes.  An- 
terior third  of  carapace  white,  median 
white  line  reaching  thoracic  furrow  (Fig. 
65).  Proximal  halves  of  femora  white,  re- 
mainder black  with  distal  white  marks  on 
dorsal  surfaces.  Patellae  inostly  black,  re- 


maining articles  white  with  black  marks  at 
base  of  setae.  Femur  I  with  row  of  three 
to  five  macrosetae  on  anterior  side;  three 
to  four  on  anteroventral  side.  Anterior 
margin  of  dorsal  abdomen  black;  dorsal  fo- 
lium yellowish  with  brown  speckles,  white 
fleur-de-lis  pattern  with  wide  branches 
(Fig.  65).  Venter  black  with  wide  white 
median  line,  flanked  by  pair  of  thin  white 
lines;  pair  of  white  spots  on  either  side  of 
spiracle  connected  by  white  line  (Fig.  66). 
Sternum  black  with  posterior  white  mark 
(Fig.  66).  Batio  of  eye  diameters:  posterior 
medians  and  anterior  medians  1.0,  anterior 
laterals  1.1,  posterior  laterals  1.1.  Anterior 
median  eyes  separated  by  1.7  diameters, 
posterior  median  eyes  by  1.0,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  3.3  diameters  of  anterior  lateral  eyes, 
lateral  eyes  separated  by  0.1  their  diame- 
ters. Total  length  8.4  mm.  Carapace  3.7 
mm  long,  2.9  wide.  First  femur  4.2  mm, 
patella  and  tibia  4.1,  metatarsus  3.7,  tarsus 
1.2.  Second  patella  and  tibia  3.4  mm,  third 

2.1,  fourth  3.1. 

Male  from  Angol,  Malleco,  Chile.  Car- 
apace reddish  brown,  anterior  third  white, 
median  white  line  extending  to  thoracic 
furrow  (Fig.  63).  Proximal  halves  of  fem- 
ora, white,  distal  halves  black.  Patellae 
black,  remaining  articles  white  widi  black 
spots  at  base  of  setae.  Feinur  I  with  row 
of  six  to  eight  macrosetae  on  anterior  side; 
seven  to  11  on  anteroventral  side.  Dorsal 
abdomen  white,  marbled,  and  speckled 
brown  (Fig.  63).  Venter  dark  brown  with 
wide  white  median  mark,  flanked  by  pair 
of  thin  white  lines;  pair  of  white  spots  on 
either  side  of  spiracle  connected  by  white 
line  (Fig.  64).  Sternum  dark  brown  with 
partly  broken  median  white  line  (Fig.  64). 
Batio  of  eye  diameters:  posterior  medians 
and  anterior  medians  1.0,  anterior  laterals 

1.2,  posterior  laterals  1.1.  Anterior  median 
eyes  separated  by  1.7  diameters,  posterior 
median  eyes  by  1.1,  anterior  median  eyes 
separated  from  anterior  laterals  by  2.9  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.2  their  diameters.  To- 
tal length  6.1  mm.  Carapace  3.2  mm  long. 


Metepeira  •  Piel 


2.5  wide.  First  femur  5.2  mm,  patella  and 
tibia  5,  metatarsus  4.9,  tarsus  1.2.  Second 
patella  and  tibia  4.1  mm,  third  2,  fourth 
3.1. 

Diagnosis.  Female  M.  rectangula  differ 
from  the  other  species  in  the  M.  vigilax 
group  (M.  vigilax,  M.  glomerahilis,  M.  ca- 
jahamba)  by  the  shape  of  the  epigynal 
openings:  from  a  ventral  view  they  are 
larger,  wider,  and  more  gaping  than  in  the 
other  species  (compare  Fig.  62  with  Figs. 
40,  48,  55).  Males  differ  from  other  spe- 
cies in  the  group  by  having  an  embolus 
that  is  relatively  larger  than  that  of  M. 
glomerahilis  and  M.  cajabaniba,  yet  small- 
er than  that  of  M.  vigilax  (compare  Fig.  60 
with  Figs.  38,  46,  53).  Among  all  members 
of  the  M.  vigilax  group,  the  keel  on  the 
median  apophysis  of  M.  rectangula  is  the 
largest  and  most  robust,  and  the  dorsal  fo- 
lium has  the  lightest  coloration  (Fig.  65). 

Va nation.  Average  body  length  of  11  fe- 
males examined  8.6  mm,  range  5.8  to  10 
mm.  Average  body  length  of  six  males  ex- 
amined 6  mm,  range  4.1  to  7  mm.  Speci- 
mens from  two  localities  in  western  Ar- 
gentina resemble  M.  vigilax,  and  may  be 
hybrids. 

Natural  History.  This  species  appears  to 
follow  a  narrow  ecological  zone  that  de- 
creases in  elevation  with  increasing  south- 
ern latitude  (Fig.  36).  Median  elevation, 
about  500  m.  Mature  specimens  have 
been  collected  January  through  April  (Fig. 
306).  Specimens  from  localities  south  of 
the  36th  parallel  tend  to  be  found  in  Jan- 
uaiy  and  Februaiy,  whereas  those  north  of 
the  36th  parallel  tend  to  be  found  in 
March  and  April. 

Distribution.  Chilean  Andes  between 
31°  and  38°  south  (Map  4). 

Records  Examined.  ARGENTINA  Cordoba:  Cala- 
muchita,  32°4'S,  64°33'W,  15.iii.l954  (J.  M.  Viana, 
MACN).  Mendoza:  Mendoza,  32°53'S,  68°49'W, 
30.iii.l965  (H.  W.  Levi,  MCZ).  CHILE  Bio-Bio:  4  km 
E  road  to  Pinto,  36°42'S,  71°53'W,  4.i.l976  (B.  Mo- 
reno, AMNH);  Road  to  Pemuco,  Ci-uce  del  Carmen, 
36°56'S,  72°4'W,  10.i.l976  (G.  Moreno,  AMNH).  Co- 
qnimbo:  lUapel:  Salamanca:  Fundo  Tahuinco, 
31°44'S,  71°5'W,  30.iv.l946  (R.  Doneso,  AMNH). 
Malleco:   Angol,   37°48'S,   72°43'W   (D.    S.    Bullock, 


CAS),  10.iii.l945  (E.  A.  Chapin,  USNM).  Maule:  10 
km  S  Curico,  35°4'S,  71°14'W,  15.iii.l968  (L.  E. 
Peiia,  MCZ);  Cordillera  de  Parral  [?],  36°9'S, 
71°50'W,  25.ii.1956  (L.  E.  Pena,  IRSNB);  Linares, 
35°51'S,  71°36'W  (L.  E.  Peiia,  IRSNB);  Miraflores, 
Pedag.  [?].  35°55'S,  71°39'W  (Toro,  AMNH). 
O'Higgins:  Fundo  Millahue,  Cunaco,  34°36'S, 
71°16'W,  30.iv.l961  (AMNH).  Region  Metropolitana: 
Melipilla,  33°42'S,  71°13'W  (L.  E.  Pena,  IRSNB). 
Valparaiso:  Casablanca,  33°19'S,  71°25'W,  15.ii.l955 
(Edwdn  Reed,  AMNH). 


Metepeira  labyrinthea  Group 

Levi  (1977)  described  the  M.  labyrin- 
thea group  veiy  broadly — it  included  spe- 
cies with  a  longitudinal  white  line  down 
the  sternum  and  a  short  keel  on  the  me- 
dian apophysis.  Here,  this  species  group  is 
much  narrowed  to  include  only  three 
North  American  species:  Metepeira  laby- 
rinthea, Metepeira  lacandon,  and  Metepei- 
ra spinipes.  Males  of  these  three  species 
are  unique  among  Metepeira  by  having  a 
toothless,  smooth  keel  on  the  median 
apophysis.  In  addition,  their  distal  embolic 
apophysis  rises  away  (anteriorly)  from  the 
embolus  proper  and  projects  forward  (ven- 
trally)  until  it  is  almost  even  with  the  em- 
bolus tip  (Figs.  67,  76).  In  contrast,  other 
Metepeira  species  with  distal  embolic 
apophyses  have  the  embolus  tip  extend  far 
beyond  the  projection  of  the  apophysis 
(e.g.,  Fig.  171).  The  female  epigynum  has 
a  characteristic  shape.  The  scape  is  thick 
and  fleshy  and  the  epigynal  openings  have 
membranes  that  make  them  look  distinctly 
(Fig.  69)  or  indistinctly  (Fig.  78)  almond- 
shaped.  The  epigyna  of  the  M.  labyrinthea 
species  group  are  easily  confused  with  that 
of  the  closely  related  species,  M.  gosoga 
(Fig.  173).  Although  most  differences  be- 
tween M.  labyinthea  group  and  M.  go,soga 
are  only  obvious  in  the  epigynum,  it 
should  be  noted  that  the  dark  marks  inside 
the  epigynal  openings  of  the  former  ap- 
pear to  look  cross-eyed,  but  this  cannot  be 
said  for  the  latter.  In  this  work,  only  the 
species  collected  at  localities  south  of  the 
U.S. /Mexico  boarder  are  treated. 


34         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


9.  Metepeira  spinipes 
F.  O.  P. -Cambridge 
Figures  67-75,  335;  Map  6 

Metepeira  spinipes  F.  O.  P.-Cambridge,  1903:  459, 
figs.  9,  10,  (J,  ?.  Male  holotype  from  Mexico  City, 
Mexico,  in  BMNH,  examined.  Roewer,  1942:  868. 

Epeira  labyrinthea  grinnelli  Coolidge,  1910:  281,  9. 
Holotype  from  Palo  Alto,  California,  lost. 

Araneus  spinipes: — Petrunkevitch,  1911:  317. 

Aranea  labyrinthea  grinnelli: — Moles,  1921:  42. 

Metepeira  douglasi  Chamberlin  and  Ivie,  1941:  18, 
figs.  21-23,  9 .  Female  holotype  from  Santa  Ana, 
California,  in  AMNH,  examined.  Chamberlin  and 
Ivie,  1942:  66,  figs.  169-170.  First  synonymized 
with  M.  labyrinthea  grinnelli  by  Levi,  1977:  198. 

Metepeira  labyrinthea  grinnelli: — Roewer,  1942:  868. 

Metepeira  labyrinthea: — Bonnet,  1957:  2822.  Erro- 
neous synonyiTiy. 

Metepeira  grinnelli: — Levi,  1977:  198,  figs.  21-27,  S , 
? .  NEW  SYNONYMY. 

Description.  Female  from  Huitzilac, 
Morelos,  Mexico.  Brown  carapace  with  an- 
terior portion  darker  reddish  brown,  white 
behind  lateral  eyes  (Fig.  74).  Legs  yellow- 
ish, femora  reddish  brown  distally,  otlier 
articles  dark  brown  distally.  Femur  I  with 
row  of  four  to  five  macrosetae  on  anterior 
side;  two  to  seven  on  anteroventral  side. 
Anterior  shoulders  of  abdomen  black. 
Dorsal  folium  with  usual  Metepeira  pat- 
tern, though  largest  branches  of  white 
fleur-de-lis  shape  usually  widened  into 
large  spots  (Fig.  74).  Venter  of  abdomen 
black  with  wide  white  median  line  (Fig. 
75).  Pair  of  small  white  spots  on  either  side 
of  spiracle.  Sternum  black  with  posterior 
white  mark  that  in  some  cases  extends  an- 
teriorly to  the  labium  (Figs.  73,  75).  Ratio 
of  eye  diameters:  posterior  medians  and 
anterior  medians  1.1,  anterior  laterals  1.0, 
posterior  laterals  0.9.  Anterior  median 
eyes  separated  by  1.9  diameters,  posterior 
median  eyes  by  1.3,  anterior  median  eyes 
separated  from  anterior  laterals  by  2.9  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.2  their  diameters.  To- 
tal length  9.2  mm.  Carapace  3.7  mm  long, 
2.9  wide.  First  femur  4.4  mm,  patella  and 
tibia  4.7,  metatarsus  4.6,  tarsus  1.3.  Sec- 
ond patella  and  tibia  4.1  mm,  third  2.4, 
fourth  3.6. 

Male  from  Huitzilac,  Morelos,  Mexico. 


Carapace,  dorsum,  venter,  sternum  as  in 
female  (Figs.  71,  72).  Distal  portions  of  leg 
articles  reddish  black,  elsewhere  yellowish. 
Femur  I  with  row  of  four  to  six  macrosetae 
on  anterior  side;  five  to  nine  on  anterov- 
entral side.  Ratio  of  eye  diameters:  poste- 
rior medians  and  anterior  medians  1.1,  an- 
terior laterals  1.2,  posterior  laterals  1.1. 
Anterior  median  eyes  separated  by  1.8  di- 
ameters, posterior  median  eyes  by  1.1,  an- 
terior median  eyes  separated  from  anterior 
laterals  by  3.2  diameters  of  anterior  lateral 
eyes,  lateral  eyes  separated  by  0.3  their  di- 
ameters. Total  length  7.5  mm.  Carapace 
3.8  mm  long,  2.8  wide.  First  femur  6.2 
mm,  patella  and  tibia  6.3,  metatarsus  7.1, 
tarsus  1.8.  Second  patella  and  tibia  5.3 
mm,  third  2.5,  fourth  4. 

Diagnosis.  Inside  each  epigynal  depres- 
sion of  M.  spinipes  and  M.  lacandon  is  a 
membrane  that  forms  a  slanted,  oval- 
shaped  opening  (Figs.  69,  78).  Within  each 
oval-shaped  opening  is  a  dark  mark,  which 
in  M.  spinipes  takes  up  a  small  part  of  that 
opening,  resulting  in  a  cross-eyed  appear- 
ance (Fig.  69).  Also,  in  M.  spinipes  (Fig. 
69),  the  edges  of  the  epigynal  depressions 
are  less  distinct  than  in  M.  lacandon  (Fig. 
78).  The  distal  embolic  apophysis  of  M. 
spinipes  is  arrow-shaped  (Fig.  79),  in  con- 
trast to  a  wider  shovel-shape  in  M.  lacanA 
don  (Fig.  79). 

Variation.  Average  body  length  of  54  fe- 
males examined  9.1  mm,  range  5.5  to  12.4 
mm.  Average  body  length  of  55  males  ex- 
amined 6.6  mm,  range  3  to  10.8  mm. 

Natural  History.  Mature  specimens 
have  been  collected  between  August  and 
early  November  (Fig.  335).  Elevations 
range  from  near  sea  level  in  California  to 
2,600  m  in  central  Mexico.  Variation  in 
sexual  dimoi-phism  appears  to  correlate 
with  habitat  and  social  structure  (Piel, 
1996).  Webs  are  found  in  dry  regions 
among  mesquite,  Opuntia,  Agave  (ma- 
guey), cultivated  Yucca,  and  Cactus.  Spi- 
ders live  in  medium  to  small  social  colo- 
nies, which  vary  in  size  in  accordance  with 
local  habitat  quality  (Uetz,  1988a,b).  This 
behavioral   and   ecological   relationship  is 


Metepeira  •  Piel        35 


4  M 


82 

lacandon 
(10) 


Figures  60-66.  Metepeira  rectangula  (Nicolet)  (sp.  8;  37°48'S,  72°43'W).  60,  male  palpus,  mesal.  61,  epigynum,  posterior.  62, 

epigynum,  ventral.  63,  male,  dorsal.  64,  male,  ventral.  65,  female,  dorsal.  66,  female,  ventral. 

Figures  67-75.  Metepeira  spinipes  F.  O.  P. -Cambridge  (sp.  9  [67-72,74,75]  19'=0'29"N,  99°15'50"W;  [73]  39°18'N,  123°48'W). 

67,  male  palpus,  mesal.  68,  epigynum,  posterior.  69,  epigynum,  ventral.  70,  male  embolic  division,  ventral.  71,  male,  dorsal.  72, 

male,  ventral.  73,  female,  ventral.  74,  female,  dorsal.  75,  female,  ventral. 

Figures  76-83.  Metepeira  lacandon  new  species  (sp.  10;  16°45'N,  92°38'W).  76,  male  palpus,  mesal.  77,  epigynum,  posterior. 

78,  epigynum,  ventral.  79,  male  embolic  division,  ventral.  80,  male,  dorsal.  81,  male,  ventral.  82,  female,  dorsal.  83,  female, 

ventral. 

Scale  bars:  dorsum  and  venter  figures  1.0  mm. 


36         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


thought  to  occur  as  a  result  of  the  spiders 
pursuing  a  risk-sensitive  foraging  strategy 
(Uetz,  1996). 

Distribution.  Oregon  to  central  Mexico 
(Levi,  1977,  map  1;  Map  6). 

Records  Examined.  MEXICO  Agiiascalientes:  Hwy 
45,  5.3  mi  N  Aguascalientes,  21°57'N,  102°17'W, 
7.ix.l967  (R.  E.  Leech,  REL).  Baja  California  Sun 
Sierra  Laguna,  17  air  mi  ENE  Todos  Santos, 
23°34'N,  110°0'W,  15.xii.l979  (C.  E.  Griswold,  CAS). 
Chihuahua:  22.4  mi  S  Miiaaca,  28°24'N,  107°26'W, 
23.viii.1950  (R.  Smith,  AMNH).  Durango:  10  mi  E 
El  Salto,  23°12'N,  105°52'W,  8.viii.l947  (W.  J. 
Gertsch,  AMNH);  11  km  W  Suehil,  23°35'N, 
104°5'W,  5.ix.l984  (W.  J.  Pulawsld,  CAS);  6  mi  NE 
El  Salto,  23°19'N,  105°50'W,  ll.viii.l947  (W.  J. 
Gertsch,  AMNH);  Las  Puentes  [?],  26°49'N, 
106°2'W,  23.vii.1947  (W.  J.  Gertsch,  AMNH);  Oti- 
napa,  24°11'N,  105°2'W,  12.viii.l947  (W.  J.  Gertsch, 
AMNH);  Palos  Colorados,  24°2'N,  104°54'W, 
5.viii.l947  (W.  J.  Gertsch,  AMNH);  Providencia. 
26°44'N,  105°56'W,  24.viii.1947  (A.  M.  Davis, 
AMNH);  SW  Durango,  23°59'17"N,  104°45'47'W, 
22.X.1994  (W.  H.  Piel,  MCZ).  Guanajuato:  30  mi  SE 
Leon,  6  mi  SE  Silao,  20°52'N,  101°21'W,  6.ix.l964 
(Jean  &  Wilton  Ivie,  AMNH);  6.3  mi  NW  Leon, 
21°13'N,  101°43'W,  6.ix.l967  (R.  E.  Leech,  REL); 
between  Moroleon  &  Cuitzeo,  20°5'36"N, 
101°9'28'W,  20.X.1994  (W.  H.  Piel,  MCZ);  near  San 
Miguel  de  Allende,  20°55'N,  100°45'W,  16.lx.1976 
(C.  E.  Grisv^^old,  CAS);  S  San  Miguel  de  Allende, 
20°46'36"N,  100°47'28"W,  20.x.  1994  (W.  H.  Piel, 
MCZ);  San  Miguel  de  Allende.  Road  SW  town, 
20°52'N,  100°56'W  25.x.  1982  (George  Uetz,  MCZ). 
Hidalgo:  18  mi  E  Huichapan,  off  Hwy  45,  20°23'N, 
99°22'W,  25.viii.1984  (W  D.  Sissom,  C.  Myers  &  L. 
Bom,  MCZ);  4  mi  N  Tizayuca,  19°54'N,  98°59'W 
20..xi.l946  (E.  S.  Ross,  CAS);  41  km  N  Zimapan, 
20°54'N,  99°13'W,  10.viii.l991  (W  H.  Piel  &  G.  S. 
Bodner,  MCZ);  Apulco,  20°19'N,  98°20'W,  6.X.1947 
(H.  Wagner,  AMNH);  Ozumbilla,  20°9'N,  101°16'W, 
2.X.1957  (R.  Dreisbach,  MCZ);  Pachuca,  20°7'N, 
98°44'W,  30.viii.l957  (R.  Dreisbach,  MCZ);  Tenango 
de  Doria,  20°19'N,  98°13'W,  5.x.  1947  (H.  Wagner, 
AMNH).  Jalisco:  12  mi  S  Mazamitla,  19°47'N, 
103°8'W,  5.xii.l948  (H.  B.  Leech,  CAS);  Charco 
Ondo,  30  km  W  Ojuelos,  21°47'N,  101°53'W 
25. ix. 1945  (H.  Wagner,  AMNH).  Mexico:  Ixtapan  de 
la  Sal,  18°50'N,  99°41'W,  24.viu.1946  (H.  Wagner, 
AMNH);  Nevado  de  Toluca,  19°18'N,  99°44'W, 
8.iv.l979  (George  Uetz,  MCZ);  San  Juan  Teotihu- 
acan,  19°41'N,  98°52'W,  4..xi.l939  (C.  M.  Bogert  & 
H.  E.  Vokes,  AMNH);  Tenancingo,  18°58'N, 
99°36'W,  6.LX.1946  (H.  Wagner,  AMNH),  l.x.1946 
(H.  Wagner,  AMNH),  15.x.  1946  (H.  Wagner, 
AMNH);  Tenango  del  Valle,  19°7'N,  '99°33'W, 
25.viii.1946  (H.  Wagner,  AMNH),  27.viii.1946  (H. 
Wagner,  AMNH);  Teotihuacan,  19°41'N,  98°52'W 
31.viii.l959    (A.    F.    Ai-cher,    AMNH);    Tepotzotlan, 


19°43'N,  99°13'W,  26.X.1982  (George  Uetz,  MCZ), 
21. ii.  1983  (George  Uetz,  MCZ),  5.x.  1983  (George 
Uetz,  MCZ);  Toluca,  19°18'N,  99°44'W,  10.viii.l978 
(George  Uetz,  MCZ),  l.viii.l986  (George  Uetz, 
MCZ);  Toluca,  at  bottom  of  mountain  near  Parque 
Cierra  Morelos,  19°18'N,  99°44'W,  10.viii.l978 
(George  Uetz,  MCZ);  Toluca,  E  of  town  on  Paseo 
ToUocan  [?],  19°18'N,  99°42'W,  23.X.1982  (George 
Uetz,  MCZ).  Mexico  D.  F.:  19°25'N,  99°10'W, 
12.X.1940  (H.  Wagner,  AMNH),  28.xii.1940  (R.  H. 
Crandall,  AMNH),  15.ix.l943  (H.  &  D.  Frizzell, 
AMNH);  Contreras,  19°18'N,  99°17'W,  4.xii.l944  (H. 
Wagner,  AMNH),  15.ix.l965  (N.  L.  H.  Krauss, 
AMNH);  Delegacion  Tlalpan,  Colonia  Santa  Ursula 
Xitla,  19°16'0"N,  99°10'25'W,  12.x.  1994  (W  H.  Piel, 
MCZ);  Desierto  de  los  Leones,  19°22'N,  99°16'W, 
15.ix.l941  (H.  Wagner,  AMNH);  El  Xitle,  18°61'N, 
99°17'W  [?],  12.viii.l942  (C.  Tellez,  AMNH);  Haci- 
enda Cordoba,  19°26'N,  99°10'W  [?],  29.X.1944  (H. 
Wagner,  AMNH);  Ouieros,  18°62'N,  99°17'W  [?], 
5.vii.l943  (M.  Cardenas,  AMNH);  Mexico  City, 
19°25'N,  99°10'W,  l.xi.l941  (C.  Velo,  AMNH), 
25.Lx.1957  (R.  Dreisbach,  MCZ);  Mixcoac,  19°23'N, 
99°11'W  (AMNH),  13.X.1940  (A.  F.  Archer,  AMNH); 
Mixenac,  19°25'N,  99°10'W  13.X.1940  (H.  Wagner, 
AMNH);  Pedregales,  18°60'N,  99°17'W  [?], 
15.viii.l909  (AMNH);  Petregal  [?],  18°60'N, 
99°17'W  l.xii.l943  (AMNH);  Rancho  Cordoba, 
19°27'N,  99°10'W,  29.X.1944  (H.  Wagner,  AMNH); 
Tlaplan,  19°17'N,  99°10'W,  7.viii.l991  (W.  H.  Piel  & 
G.  S.  Bodner,  MCZ).  Michoacan:  25  mi  W  La  Barca 
nr  Lago  de  Chapala,  20°17'N,  102°34'W,  ll.ix.l976 
(C.  E.  Griswold  &  Jackson,  CAS);  between  Patzcuaro 
&  Uruapan,  19°29'19"N,  101°48'20"W,  19.X.1994  (W 
H.  Piel,  MCZ);  Hills  N  of  Patzcuaro,  19°45'N, 
101°36'W,  24.viii.1959  (A.  F.  Archer,  AMNH);  Hwy 
110,  4  mi  W.  Jiquilpan,  19°59'N,  102°47'W, 
2.viii.l967  (R.  E.  Leech,  REL);  Hwy  15,  9.5  mi  W 
Morelia,  19°42'N,  101°16'W;  18.viii.l967  (R.  E. 
Leech,  REL);  Lake  Chapala,  NW  of  Cojumatlan, 
20°10'N,  102°53'W,  7.ix.l966  (Jean  &  Wilton  Ivie, 
AMNH);  Monte  de  Zacapu,  19°47'N,  101°50'W, 
24.viii.1959  (A.  F.  Archer,  AMNH).  Morelos:  Cuer- 
navaca,  18°55'N,  99°15'W  (AMNH),  15.ix.l941  (H. 
Wagner,  AMNH),  18.xi.l946  (M.  G.  Bradt,  AMNH); 
Cueniavaca/Tepotzotlan,  interchange  between  1-95  & 
115,  18°55'N,  99°13'W,  7.viii.l978  (MCZ);  Huitzilac, 
19°2'2"N,  99°16'13'W^,  13.X.1994  (W.  H.  Piel,  MCZ); 
North  of  Cuernavaca,  18°58'11"N,  99°14'37"W, 
11.x.  1994  (W.  H.  Piel,  MCZ);  S  of  Huitzilac, 
19°0'29"N,  99°15'50"\'V,  16.x.  1994  (W  H.  Piel,  MCZ). 
Puehla:  6  mi  E  Rio  Frio,  19°20'N,  98°35'W, 
22.viii.1964  (Jean  &  Wilton  Ivie,  AMNH);  Puebla, 
19°3'N,  98°12'W,  21.x.  1982  (George  Uetz,  MCZ). 
San  Luis  Potosi:  3  km  W  Pilares,  21°55'34"N, 
100°48'6'W,  21.X.1994  (W.  H.  Piel,  MCZ);  Cuidaddel 
Maiz,  22°24'N,  99°36'W,  25.viii.1954  (R.  Dreisbach, 
MCZ).  Sonora:  46  mi  S  Agua  Prieta  on  Highway  10, 
31°0'N,  109°16'W,  15.viii.l959  (B.  A.  Branson, 
AMNH);  Hermosillo,  29°4'N,  110°55'W,  20.lx.1952 
(B.  Malldn  &  V.  E.  Thatcher,  AMNH);  Sierra  Man- 


Metepeira  •  Piel 


37 


zanal,  30°50'N,  110°10'W,  14.ix.l976  (Roth  & 
Schroepfer,  MCZ).  Tlaxcala:  Huamantla,  19°19'N, 
97°56'W  [?],  15.vii.l981  (C.  Gold,  CAS).  Veracruz: 
15  mi  W.  Banderilla,  19°39'N,  97°8'W,  31.X.1973  (S. 
C.  Williams  &  C.  L.  Mullinex,  CAS);  15  mi  West  of 
Jalapa,  19°32'N,  97°9'W,  23.vi.1946  (A.  M.  &  L.  I. 
Davis,  AMNH).  Zacatecas:  13  mi  N.  Sombrerete, 
23°44'32"N,  103°47'10"W  22.X.1994  (W  H.  Piel, 
MCZ);  Canutillo,  24°47'N,  101°31'W,  14.viii.l947  (W. 
J.  Gertsch,  AMNH);  S.  Zacatecas,  22°45'7"N, 
102°29'37"W,  22.X.1994  (W.  H.  Piel,  MCZ).  USA.  Ar- 
izona: Southwestern  Research  Station,  Chiricaliua 
Mtns.,  31°35'N,  109°14'W,  20.viii.l976  (V.  Roth, 
MCZ).  California:  26  mi  W.  Santa  Rosa  on  Hwy  116, 
38°31'N,  123°4'W,  19.ix.l976  (M.  E.  Thompson, 
MCZ);  Mendocino,  39°18'N,  123°48'W,  18.viii.l959 
(W  J.  Gertsch,  MCZ);  Monterey,  36°36'N,  121°54'W, 
l.ix.l949  (A.  F.  Archer,  MCZ);  Pacific  Grove, 
36°37'N,  121°56'W  (R.  V.  Chamberlin,  MCZ);  Palo 
Alto,  37°27'N,  122°9'W  (Doane,  MCZ);  Salt  marsh 
on  N  shore  of  San  Pablo  Bay,  Vallejo,  38°8'N, 
122°27'W  (D.  Spiller,  MCZ). 

10.  Metepeira  lacandon  new  species 
Figures  76-83,  332;  Map  6 

Holotijpe.  Male  from  San  Cristobal,  Chiapas,  Mexico. 
The  specific  name  is  a  noun  in  apposition  after  the 
Indian  people  who  live  in  Chiapas.  Holotype  de- 
posited in  the  AMNH. 

Description.  Female  paratype  from  San 
Cristobal,  Chiapas,  Mexico.  Reddish  cara- 
pace, slightly  darker  in  anterior  half,  ligh- 
ter behind  lateral  eyes  (Fig.  82).  Leg  ar- 
ticles yellowish,  gradually  turning  reddish 
brown  distally.  Femur  I  with  row  of  four 
or  five  macrosetae  on  anterior  side;  two  to 
four  setae  on  anteroventral  side.  Anterior 
shoulders  of  abdomen  black.  Branches  of 
white  fleur-de-lis  shape  in  dorsal  folium 
thinner  than  in  most  species  (Fig.  82). 
Venter  of  abdomen  black  with  wide  white 
median  line  that  extends  about  half  the 
distance  between  epigynal  groove  and 
spinnerets  (Fig.  83).  Pair  of  very  small 
white  spots  on  either  side  of  spiracle.  Ster- 
num black,  often  with  central  w^hite  spot 
(Fig.  83).  Ratio  of  eye  diameters:  posterior 
medians  and  anterior  medians  1.0,  anterior 
laterals  1.3,  posterior  laterals  1.2.  Anterior 
median  eyes  separated  by  1.7  diameters, 
posterior  median  eyes  by  1.0,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  4.5  diameters  of  anterior  lateral  eyes, 
lateral  eyes  separated  by  0.1  their  diame- 


ters. Total  length  11.5  mm.  Carapace  4.6 
mm  long,  3.6  wide.  First  femur  5.3  mm, 
patella  and  tibia  5.3,  metatarsus  4.9,  tarsus 

1.6.  Second  patella  and  tibia  4.4  mm,  third 

2.7,  fourth  4. 

Male  holotype.  Carapace,  dorsum,  ven- 
ter, sternum  as  in  female  (Figs.  80,  81). 
Distal  halves  of  femora,  tibia  reddish 
brown,  elsewhere  yellowish.  Patellae, 
metatarsi  reddish.  Femur  I  with  row  of 
four  or  five  macrosetae  on  anterior  side; 
six  or  seven  on  anteroventral  side.  Ratio  of 
eye  diameters:  posterior  medians  and  an- 
terior medians  1.1,  anterior  laterals  1.5, 
posterior  laterals  1.3.  Anterior  median 
eyes  separated  by  1.7  diameters,  posterior 
median  eyes  by  0.9,  anterior  median  eyes 
separated  from  anterior  laterals  by  3.3  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.3  their  diameters.  To- 
tal length  6.8  mm.  Carapace  3.5  mm  long, 
2.7  wide.  First  feinur  5.3  mm,  patella  and 
tibia  3.9,  metatarsus  5.6,  tarsus  1.7.  Sec- 
ond patella  and  tibia  4.4  mm,  third  2.3, 
fourth  3.4. 

Diagnosis.  Inside  each  epigynal  depres- 
sion of  M.  lacandon  and  M.  spinipes  is  a 
membrane  that  forms  a  slanted,  oval- 
shaped  opening  (Figs.  69,  78).  Within  each 
oval-shaped  opening  is  a  dark  mark,  which 
in  M.  lacandon  takes  up  only  a  large  part 
of  that  opening,  resulting  in  a  less  cross- 
eyed appearance  (Fig.  78).  Also,  in  M.  la- 
candon (Fig.  78),  the  edges  of  the  epigynal 
depressions  are  luore  distinct  than  in  M. 
spinipes  (Fig.  69).  The  distal  embolic 
apophysis  of  M.  lacandon  is  more  shovel- 
shaped  (Fig.  79)  than  the  thinner,  arrow- 
shaped  one  in  M.  .spinipes  (Fig.  70). 

Variation.  Average  body  length  of  three 
females  examined  8.5  mm,  range  7.5  to  10 
mm.  Average  body  length  of  four  males 
examined  5.8  mm,  range  4.2  to  6.8  mm. 

Natural  History.  Mature  specirnens 
were  collected  between  July  and  Septem- 
ber (Fig.  332)  from  oak— pine  woodland. 
Elevations  range  from  1,700  to  2,300  m. 

Distribution.  Mountainous  regions  of 
Chiapas,  Mexico  (Map  6). 


38         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


Records  Examined.  MEXICO  Chiapas:  12  km  NW 
Comitan,  16°23'N,  92°15'W,  30.\dii.l976  (E.  S.  Ross, 
CAS);  4  mi  SE  San  Cristobal,  16°42'N,  92°36'W, 
23.viii.1966  (Jean  &  Wilton  Ivie,  AMNH);  5  km  W 
San  Cristobal  de  Las  Casas  on  HWY  190,  16°44'N, 
92°41'W,  27.vii.1983  (W.  Maddison  &  R.  S.  Anderson, 
MCZ);  5  mi  W  San  Cristobal,  16°45'N,  92°41'W, 
24.viii.1966  (Jean  &  Wilton  Ivie,  AMNH);  San  Cris- 
tobal, 16°45'N,  92°38'W,  13.ix.l947  (H.  Wagner, 
AMNH);  San  Cristobal  de  las  Casas,  16°45'N, 
92°38'W,  22.vii.1947  (C.  J.  &  M.  Goodnight, 
AMNH);  Tenejapa,  16°49'N,  92°31'W,  22.vii.1950  (C. 
J.  &  M.  Goodnight,  AMNH). 

Metepeira  nigriventris  Group 

There  are  five  species  in  the  Metepeira 
nigriventris  group:  Metepeira  nigriventris, 
Metepeira  tarapaca,  Metepeira  calamuchi- 
ta,  Metepeira  galatheae,  and  Metepeira 
karkii.  These  closely  related  species  are  of- 
ten hard  to  distinguish  because  their  gen- 
italia are  similar,  yet  highly  variable  within 
a  species.  This  species  group  is  easily  rec- 
ognized by  the  distinctive  shape  of  the 
scape  and  similarities  in  palp  moiphology. 
Typically  the  base  of  the  scape  originates 
anteriorly  and  projects  ventrally  before 
curving  posteriorly.  This  projection  creates 
an  overhang  and  a  noticeable  gap  between 
the  scape  and  the  genital  openings  (e.g., 
Figs.  85,  86,  101,  102).  The  embolus  is 
thick  and  has  a  large,  prominent,  distal 
apophysis  that  hides  under  the  terminal 
apophysis  (Figs.  84,  92,  100,  110,  121). 

1 1 .  Metepeira  nigriventris  (Taczanowski) 
Figures,  84-91,  310;  Map  5 

Epeira  nigriventris  Taczanowski,  1878:  151,  fig.  6,  ?. 
Female  lectotype  from  Lake  Junin,  Peru,  in  PAN, 
type  lost.  Keyserling,  1893:  217,  fig.  161,  9,3. 

Araneus  nigriventris: — Chamberlin,  1916:  248.  Bon- 
net, 1955:  550. 

Metepeira  nigriventris: — Chamberlin  and  Ivie,  1942: 
74,  figs.  211-214,  9,3.  Platnick,  1993:  449. 

Note.  Although  the  type  is  lost,  the  type  locality 
and  Taczanowski  s  descriptions  are  sufficient  to  rec- 
ognize the  species. 

Description.  Female  from  12  km  west  of 
Tarma,  Junin,  Peru.  Carapace  dark  brown, 
light  around  eyes  with  lateral  posterior  ex- 
tensions (Fig.  90).  Proximal  halves  of  leg 
articles  yellow,  distal  halves  black.  Femur 
I  with  row  of  four  macrosetae  on  anterior 


side;  five  on  anteroventral  side.  Dorsum 
darker  and  white  fleur-de-lis  pattern  small- 
er than  in  most  species  (Fig.  90).  Venter 
mostly  black  with  reduced,  short,  thin, 
white  median  line  (Fig.  91).  SternuiTi  en- 
tirely black.  Ratio  of  eye  diameters:  pos- 
terior medians  and  anterior  medians  1.0, 
anterior  laterals  1.4,  posterior  laterals  1.4. 
Anterior  median  eyes  separated  by  1.8  di- 
ameters, posterior  median  eyes  by  1.2,  an- 
terior median  eyes  separated  from  anterior 
laterals  by  5  diameters  of  anterior  lateral 
eyes,  lateral  eyes  separated  by  0.4  their  di- 
ameters. Total  length  11  mm.  Carapace  5.1 
mm  long,  4.2  wide.  First  femur  5.5  mm, 
patella  and  tibia  5.8,  metatarsus  5.5,  tarsus 
1.9.  Second  patella  and  tibia  5.3  mm,  third 
3.3,  fourth  4.5. 

Male  from  same  locality  as  female.  An- 
terior margin  of  chelicerae  with  large, 
swollen  tooth  and  several  denticles.  Cara- 
pace reddish  brown  with  lighter  eye  re- 
gion, lateral  posterior  extensions,  and  long, 
thin  median  white  line  (Fig.  88).  Femur  1 
with  row  of  about  five  macrosetae  on  an- 
terior side;  nine  on  anteroventral  side. 
Coloration  of  legs,  dorsum,  venter,  and 
sternum  as  in  female  (Figs.  88,  89).  Ratio 
of  eye  diameters:  posterior  medians  and 
anterior  medians  1.1,  anterior  laterals  1.3, 
posterior  laterals  1.3.  Anterior  median 
eyes  separated  by  1.8  diameters,  posterior 
median  eyes  by  1.2,  anterior  median  eyes 
separated  from  anterior  laterals  by  3.7  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.4  their  diameters.  To- 
tal length  9  mm.  Carapace  4  mm  long,  3.2 
wide.  First  femur  5.8  mm,  patella  and  tibia 
6.2,  metatarsus  6.5,  tarsus  1.7.  Second  pa- 
tella and  tibia  4.5  mm,  third  2.8,  fourth  4. 

Diagnosis.  Metepeira  nigriventris  is  eas- 
ily distinguished  from  other  species  in  the 
M.  nigriventris  group  by  its  dark  pigmen- 
tation. As  its  name  implies,  the  sternum  is 
black  and  the  white  ventral  mark  on  the 
abdomen  is  reduced  to  a  much  shorter  and 
thinner  line  (Figs.  88-91).  While  the  ster- 
num of  M.  karkii  is  similarly  dark,  the  dor- 
sal and  ventral  markings  on  the  abdomen 
are   much   lighter   (compare   Figs.   88-91 


Metepeira  •  Piel        39 


nigriventris 
(11) 


i9&   9^^^       99 

«        tarapaca 


(12) 


A.  iBi 


H  106 107  _ 

calamuchita 
105  -  (13) 


Figures  84-91.  Metepeira  nigriventris  (Taczanowski)  (sp.  11;  ir25'S,  75°48'W).  84,  male  palpus,  mesal.  85,  epigynum,  pos- 
terior. 86,  epigynum,  ventral.  87,  male  embolic  division,  ventral.  88,  male,  dorsal.  89,  male,  ventral.  90,  female,  dorsal.  91, 

female,  ventral.  ,  „„       . 

Figures  92-99    Metepeira  tarapaca  new  species  (sp.  12;  21°39'S,  69°33'W).  92,  male  palpus,  mesal.  93,  epigynum,  postenor. 

94,  epigynum,  ventral.  95,  male  embolic  division,  ventral.  96,  male,  dorsal.  97,  male,  ventral.  98,  female,  dorsal.  99,  female, 

ventral.  ,  ,.,„.,• 

Figures  100-107.  Metepeira  calamuchita  new  species  (sp,  13;  32°4'S,  64°33'W).  100,  male  palpus,  mesal.  101,  epigynurri, 
posterior.  102,  epigynum,  ventral.  103,  male  embolic  division,  ventral.  104,  male,  dorsal.  105,  male,  ventral.  106,  female,  dorsal. 
107,  female,  ventral. 
Scale  bars:  dorsum  and  venter  figures  1.0  mm. 


40         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


with  Figs.  125-128).  The  epigynum  of  M. 
nigriventris  has  a  wider  membrane  over 
the  openings  that  is  visible  on  either  side 
of  a  very  wide  scape  (Fig.  86).  In  contrast, 
this  feature  is  hidden  behind  a  thinner 
scape  in  its  Hkely  sister  species,  M.  tara- 
paca  (Fig.  94).  The  overall  shape  of  tlie 
epigynum  (Fig.  86)  and  the  pair  of  small 
notches  in  the  posterior  lobes  (Fig.  85) 
also  make  this  species  distinctive.  The 
male  palp  of  M.  nigriventris  has  an  em- 
bolus that  is  relatively  sliinmer  and  more 
graceful  than  those  of  other  species  in  the 
M.  nigriventris  species  group  (compare 
Fig.  84  with  Figs.  92,  100,  109-110,  121). 
The  shape  of  the  embolus  and  its  distal 
apophysis  differs  from  other  species  (com- 
pare Fig.  87  with  Figs.  95,  103,  112,  124). 

Variation.  Average  body  length  of  25  fe- 
males examined  9.5  mm,  range  7.2  to  11.5 
mm.  Average  body  length  of  16  males  ex- 
amined 7.4  mm,  range  5.3  to  9.5  mm. 

Natural  History.  Spiders  are  commonly 
found  around  Lake  Titicaca  living  in  me- 
dium and  large  colonies  among  power 
lines,  Bolivian  pines.  Cactus,  Bacharis, 
rock  outcroppings,  and  tall  grasses  (L. 
Rayor,  personal  communication,  and  vari- 
ous locality  labels).  Mature  speciinens 
have  been  collected  throughout  the  year 
except  September  and  October  (Fig.  310). 
Median  elevation,  3,900  m. 

Distribution.  High  altitude  regions  of 
southern  Peru  and  western  Bolivia  (Map 
5). 

Records  Examined.  ARGENTINA  Jujuy:  Puma- 
huasi,  22°17'S,  65°41'W,  8.xi.l970  (L.  E.  Pena, 
MCZ).  BOLIVIA  La  Paz:  45  mi  S  La  Paz,  17°9'S, 
67°36'W,  25.ii.1951  (E.  S.  Ross  &  Michelbacher, 
CAS);  70  mi  S  La  Paz,  17°30'S,  67°36'W,  25.ii.1951 
(E.  S.  Ross  &  Michelbacher,  CAS);  La  Paz,  Avenida 
Sport  Club,  16°30'S,  68°9'W,  4.i.l959  (A.  M.  Nadler, 
AMNH);  La  Paz,  in  garden  of  house,  16°30'S, 
68°9'W,  15.iv.l959  (R.  Walsh,  AMNH);  Lake  Titicaca, 
Copacabana,  Yampupata,  &  Isla  del  Sol,  16°10'S, 
69°5'W,  17.V.1995  (L.  Rayor,  MCZ);  near  La  Paz, 
16°30'S,  68°9'W,  24.V.1958  (R.  Walsh,  AMNH);  S  end 
of  Lake  Titicaca,  100  km  NW  La  Paz,  16°10'S, 
69°5'W,  5.vii.l958  (R.  Walsh,  AMNH);  Tialiuanaco, 
Puma  Puerto  Ruins,  16°33'S,  68°42'W,  l.ii.l973  (Ann 
Moreton,  MCZ).  Oruro:  6  km  N  Challapata,  18°51'S, 
66°47'W,   23.ii.1951    (E.    S.    Ross    &    Michelbacher, 


CAS);  Gorge  Uhuschlucht,  near  Oruro,  17°59'S, 
67°9'W,  7.ii.l954  (Forster  &  Schindler,  ZSM).  Potosi: 
Villazon,  22°6'S,  65°36'W,  30.xii.l984  (L.  E.  Peiia, 
AMNH).  PERU  Apurimac:  Chincheros,  13°30'48"S, 
73°42'47"W,  12.xii.l980  (C.  Gold,  CAS);  Puna  near 
Abancay,  13°38'2"S,  72°52'52"W,  15.xii.l947  (W.  Wey- 
rauch,  CAS).  Ayacucho:  Puquio,  14°42'S,  74°8'W, 
15. iv.  1950  (F.  Blancas,  MUSM);  San  Antonio  (Pu- 
quio), 14°47'S,  74°7'W,  l.xi.l985  (D.  Silva,  MUSM). 
Cusco:  Cheqquerec,  13°23'S,  72°8'W,  2.ix.l993  (J. 
Ochoa  Camara,  MCZ);  Cusco,  13°31'6"S, 
71°58'41"W,  8.viii.l965  (P  &  B.  Wygodzinsky, 
AMNH).  junin:  8  mi  W  Tarma,  11°25'S,  75°48'W, 
6.i.l955  (E.  I.  Schlinger  &  E.  S.  Ross,  CAS);  Cochas 
Bajo,  11  km  W  Tarma,  11°25'21"S,  75°46'11"W, 
27iii.l988  (J.  Palmer  &  D.  Smith,  MCZ);  Cochas 
Bajo,  11  km  W  Tarma,  rock  ledge  in  agricultural  val- 
ley, 11°25'21"S,  75°46'11"W,  29.iii.1988  (J.  Palmer, 
MCZ);  Huancayo,  12°4'S,  75°14'W,  15.vi.l947  (W. 
Weyrauch,  AMNH);  Oroya,  11°32'S,  75°54'W, 
12.iv.l914  (M.  P.  Anderson,  AMNH).  Lima:  Bosque 
de  Zarate,  11°53'S,  76°27'W,  18.i.l981  (J.  Francke, 
MUSM).  Puno:  10  mi  N  Ayaviri,  14°45'S,  70°35'W, 
l.iii.l951  (E.  S.  Ross  &  Michelbacher,  CAS);  Cama- 
cane,  15°55'S,  69°50'W,  20.xi.l955  (L.  E.  Peiia, 
IRSNB);  Isla  Taquih,  Lago  Titicaca,  15°46'S, 
69°41'W,  23.xii.1980  (C.  Gold,  CAS);  Juh  (col.  Chu- 
cuito),  16°13'S,  69°27'W,  7.xi.l952  (F.  Blancas, 
MUSM);  near  Chucuito,  Lago  Titicaca,  15°50'S, 
69°48'W,  10.iii.l953  (M.  Koepcke,  MUSM);  Puna, 
Lake  Titicaca,  15°50'S,  70°2'W,  15.vi.l947  (W  Wey- 
rauch, AMNH);  Puno,  15°50'S,  70°2'W  (Soukup, 
AMNH);  Yunguyo,  downtown  plaza,  16°15'S,  69°5'W, 
31.i.l973  (Ann  Moreton,  MCZ). 

12.  Metepeira  tarapaca  new  species 
Figures  92-99,  305;  Map  7  | 

Holotype.  Male  from  Quillagua,  Antofagasta,  Chile, 
4.ii.l965,  L.  E.  Pena,  in  MCZ.  The  specific  name 
is  a  noun  in  apposition  after  a  Chilean  province 
where  it  is  abundant. 

Description.  Female  paratype  from 
Quillagua,  Antofagasta,  Chile.  Light  red- 
dish brown  carapace,  white  in  center  and 
around  eyes  with  lateral  posterior  exten- 
sions (Fig.  98).  Legs  yellowish  white, 
ringed  brown  at  distal  ends  of  articles.  Fe- 
mur I  with  row  of  four  macrosetae  on  an- 
terior side;  three  on  anteroventral  side. 
Dorsal  folium  white  with  black  speckles 
(Fig.  98).  Venter  of  abdomen  black  with 
wide,  white  median  line,  sometimes 
flanked  by  thinner  white  lines  that  togeth- 
er form  a  U-shape  posteriorly  (Fig.  98). 
Sternum  black  with  median  white  line, 
sometimes  broken  (Fig.  99).  Ratio  of  eye 


Metepeira  •  Piel        41 


4500j 
400O- 
350O- 
3000-- 
250a- 

20oa- 

1500- - 

1000-- 

500- - 

0  .- 


cP       o    ° 


0 


20  40  60  80  100 

Percent  of  median  white  line  on  sternum 


Figure  108.  Elevation  of  collection  localities  of  mature  female 
M.  tarapaca  with  differing  amounts  of  whiite  on  the  sternum. 
Spiders  with  a  median  white  line  covering  100%  of  the  sternum 
length  are  found  at  a  wide  altitude  range.  Spiders  with  a  me- 
dian white  mark  covering  only  a  short  length  of  the  sternum 
are  only  found  at  high  elevations.  Elevations  estimated  from 
NOAA  database  of  5-  by  5-minute  geographic  tiles. 


diameters:  posterior  medians  and  anterior 
medians  1.0,  anterior  laterals  1.3,  posterior 
laterals  1.2.  Anterior  median  eyes  separat- 
ed by  1.4  diameters,  posterior  median  eyes 
by  1.0,  anterior  median  eyes  separated 
from  anterior  laterals  by  3.4  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.1  their  diameters.  Total  length  7.8 
mm.  Carapace  3.2  mm  long,  2.8  wide. 
First  femur  4.3  mm,  patella  and  tibia  4.4, 
metatarsus  4,  tarsus  1.2.  Second  patella 
and  tibia  3.8  mm,  third  2.1,  fourth  3.2. 

Male  holotype.  Light  reddish  brown 
carapace,  lighter  around  eyes  and  white 
mark  in  center  (Fig.  96).  Legs  yellowish 
white,  gradually  growing  darker  toward 
distal  ends  of  articles.  Femur  I  with  row 
of  four  macrosetae  on  anterior  side;  five  on 
anteroventral  side.  Dorsal  folium,  venter, 
and  sternum  as  in  female  (Figs.  96,  97). 
Ratio  of  eye  diameters:  posterior  medians 
and  anterior  medians  1.0,  anterior  laterals 
1.2,  posterior  laterals  1.1.  Anterior  median 
eyes  separated  by  1.5  diameters,  posterior 
median  eyes  by  1.0,  anterior  median  eyes 
separated  from  anterior  laterals  by  2.5  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.1  their  diameters.  To- 
tal length  4.4  mm.  Carapace  2  mm  long, 
1.6  wide.  First  femur  3.3  mm,  patella  and 
tibia  3.5,  metatarsus  3.3,  tarsus  1.  Second 
patella  and  tibia  2.8  mm,  third  1.4,  fourth 
2.2. 

Diagnosis.  Unlike  M.  nigriventris  or  M. 


karkii,  M.  tarapaca  has  a  white  mark  on 
the  sternum  (Figs.  97,  99).  The  epigynum 
is  close  to  that  of  M.  nigriventris,  except 
that  it  is  less  robust  (compare  Fig.  93  with 
Fig.  85),  and  the  membrane  just  over  the 
openings  cannot  be  seen  behind  the  thin- 
ner scape  (compare  Fig.  86  with  Fig.  94). 
Unlike  M.  calamuchita,  the  epigynum  of 
M.  tarapaca  does  not  widen  posteriorly 
(compare  Fig.  94  with  Fig.  102);  unlike  M. 
karkii,  the  posterior  epigynal  lobes  are  not 
swollen  (compare  Fig.  94  with  Fig.  123); 
unlike  M.  galatheae,  the  black  sclerotized 
circles  behind  the  epigynal  openings  are 
larger  and  not  shifted  posteriorly  (compare 
Fig.  94  with  Figs.  118-120).  The  segment 
of  the  embolus  between  the  basal  and  dis- 
tal embolic  apophysis  is  relatively  thicker 
than  in  other  members  of  the  M.  nigriven- 
tris species  group  (compare  Fig.  92  with 
Figs.  84,  100,  109,  121).  The  terminal  di- 
vision of  the  male  palp  in  M.  tarapaca 
shows  a  distal  embolic  apophysis  that  dif- 
fers in  shape  from  that  of  other  species 
(compare  Fig.  95  with  Figs.  87,  103,  117, 
124). 

Variation.  Average  body  length  of  22  fe- 
males examined  7.4  inm,  range  5  to  11.5 
mm.  Average  body  length  of  six  males  ex- 
amined 5.9  mm,  range  4.4  to  7.3  mm.  The 
dorsum  and  venter  are  often  darker  than 
the  holotype.  Many  lack  the  flanking  lines 
and  the  U-shape  on  the  venter;  some  at 
higher  elevation  have  a  much  reduced 
white  line  on  the  sternum. 

Natural  History.  Mature  specimens 
have  been  collected  throughout  the  year, 
especially  between  January  and  April  (Fig. 
305).  V.  Roth  (vial  label)  notes  that  these 
spiders  live  in  a  social  colony.  M.  Roy  (per- 
sonal communication)  reports  that  colo- 
nies can  reach  200  individuals.  Median  el- 
evation, 2,800  m. 

Distribution.  Moderately  high  altitudes 
in  northern  Chile  and  southern  Peru  (Map 
7). 

Records  Exmnined.  CHILE  Antofagasta:  7  km  S 
Toconao,  23°11'S,  68°1'W,  25.xii.1988  (V.  &  B.  Roth, 
CAS);  Aguas  Blancas  [?]  (=  24°11'S  69°51'W),  To- 
conao,   23°11'S,    68°1'W,    11.x.  1955    (L.    E.    Peiia, 


42         Bulletin  Museum  of  Coinparative  Zoology,  Vol.  157,  No.  1 


IRSN);  Antofagasta,  23°39'S,  70°24'W,  15.xi.l975  (L. 
E.  Pena,  AMNH);  Quillagua,  21°39'S,  69°33'W, 
4.ii.l965  (L.  E.  Pena,  MCZ),  21.i.l973  (W.  C.  Sedg- 
wick, MCZ);  Rio  Loa,  25  km  S  Quillagua,  21°54'S, 
69°33'W,  20.viii.l966  (E.  Schlinger  &  M.  Irwin, 
CAS).  Elloa:  Thermo  Puritania,  35  km  N  San  Pedro 
de  Atacama,  22°37'S,  68°13'W,  25.xii.1988  (V.  &  B. 
Roth,  CAS).  Tarapaca:  Arica  Timar,  Alociado  [?], 
18°45'S,  69°42'W,  22.iii.1973  (N.  Hichins,  AMNH); 
Ariquilda,  19°38'S,  69°32'W,  29.iv.1969  (L.  E.  Pena, 
MCZ);  Canchones,  20°27'S,  69°37'W,  29.i.l973  (W. 
C.  Sedgwick,  MCZ);  Chapiquilta,  19°18'S,  69°25'W, 
6.vi.l968  (L.  E.  Peiia,  MCZ);  Chiapa,  19°32'S, 
69°13'W,  24. iv.  1969  (L.  E.  Peiia,  MCZ);  Pica, 
20°30'S,  69°21'W,  26.i.l973  (W.  C.  Sedgwick,  MCZ); 
Poroma,  Quebrada  de  Tarapaca,  19°52'S,  69°11'W, 
20.iv.l969  (L.  E.  Peiia,  MCZ);  Quisama,  19°19'S, 
69°28'W,  5.vi.l968  (L.  E.  Peiia,  MCZ).  PERU  Are- 
quipa:  Arequipa,  16°23'56"S,  71°32'6"W,  15.vii.l955 
(A.  Meza,  MZSP);  Chiguata,  near  Arequipa,  16°24'S, 
71°24'W,  15.ii.l948  (W.  Weyrauch,  MUSM). 

1 3.  Metepeira  calamuchita  new  species 
Figures  100-107,  308;  IVIap  5 

Holotype.  Male  from  Calamuchita,  Cordoba,  Argen- 
tina, 15.i.l955,  J.  M.  Viana,  in  MACN.  The  specific 
name  is  a  noun  in  apposition  after  tlie  locality. 

Description.  Female  paratype  from  Cal- 
amuchita, Cordoba,  Argentina.  Carapace 
reddish  brown,  light  around  eyes  with  lat- 
eral posterior  extensions  (Fig.  106).  Leg 
articles  annulated  distally.  Femur  I  with 
row  of  six  or  seven  macrosetae  on  anterior 
side;  five  on  anteroventral  side.  Anterior 
portion  of  dorsal  folium  lighter  than  in 
most  species  (Fig.  106).  Venter  black  with 
wide  white  median  line  and  pair  of  large 
w^hite  spots  on  either  side  of  spiracle  (Fig. 
107).  Sternum  brownish  black  with  wide, 
white  median  line  (Fig.  107).  Ratio  of  eye 
diameters:  posterior  medians  and  anterior 
medians  1.1,  anterior  laterals  1.4,  posterior 
laterals  1.3.  Anterior  median  eyes  separat- 
ed by  1.6  diameters,  posterior  median  eyes 
by  0.9,  anterior  median  eyes  separated 
from  anterior  laterals  by  3.2  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.2  their  diameters.  Total  length  8.9 
mm.  Carapace  3.4  mm  long,  2.8  wide. 
First  femur  4.5  mm,  patella  and  tibia  4.7, 
metatarsus  4.7,  tarsus  1.3.  Second  patella 
and  tibia  4.1  mm,  third  2.3,  fourth  3.5. 

Male   holotype.    Carapace   brown  with 


light,  triangular,  median  mark  pointing 
posteriorly  (Fig.  104).  Leg  articles  white, 
darkening  brown  distally.  Femur  I  with 
row  of  four  macrosetae  on  anterior  side; 
five  or  six  on  anteroventral  side.  Anterior 
dorsal  folium  mostly  white,  posterior  most- 
ly black  (Fig.  104).  Venter  as  in  female 
(Fig.  105).  Broad  white  median  mark  on 
sternum  (Fig.  105).  Ratio  of  eye  diame- 
ters: posterior  medians  and  anterior  me- 
dians 1.1,  anterior  laterals  1.4,  posterior 
laterals  1.4.  Anterior  median  eyes  separat- 
ed by  1.4  diameters,  posterior  median  eyes 
by  0.9,  anterior  median  eyes  separated 
from  anterior  laterals  by  2.2  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.2  their  diameters.  Total  length  4.4 
mm.  Carapace  2.3  mm  long,  1.8  wide. 
First  femur  3.8  mm,  patella  and  tibia  3.7, 
metatarsus  4,  tarsus  1.2.  Second  patella 
and  tibia  2.9  mm,  third  1.5,  fourth  2.5. 

Diagnosis.  Metepeira  calainuchita  dif- 
fers from  others  in  the  M.  nigriventris  spe- 
cies group  by  the  large,  scooped  openings, 
and  posteriorly  widening  epigynum  (Fig. 
102).  The  epigyna  of  M.  tarapaca  and  M. 
calamuchita  both  have  deep  scooped-out 
openings,  but  these  are  angled  posteriorly 
in  the  latter  (Fig.  102)  and  laterally  in  the 
former  (Fig.  94).  The  male  palp  of  M.  cal- 
amuchita is  easily  distinguished  from  that 
of  other  species  by  the  widened  (Fig.  100) 
and  inwardly  curved  (Fig.  103)  "dewlap" 
under  the  embolus. 

Variation.  Average  body  length  of  13  fe- 
males examined  7.7  mm,  range  5  to  9.8 
mm.  Average  body  length  of  five  males  ex- 
amined 5.2  mm,  range  4  to  6.4  mm.  Two 
specimens  collected  near  2,700  m  in  Itur- 
be,  Jujuy,  Argentina,  have  entirely  dark 
sterna.  In  contrast,  most  specimens  are 
found  at  lower  elevations  with  white  me- 
dian lines  on  their  sterna. 

Natural  Historij.  Mature  specimens 
have  been  collected  between  October  and 
July,  although  they  are  probably  available 
throughout  the  year  (Fig.  308).  Most  ele- 
vations range  from  150  to  1,700  m,  with 
one  population  at  2,700  m. 

Distribution.     Northern    Argentina,    at 


Metepeira  •  Piel        43 


mostly  lower  altitudes  east  of  the  Andes 

(Map.  5). 

Records  Examined.  ARGENTINA  Buenos  Aires: 
Las  Espaduiias  [?],  Sierra  de  la  Ventana,  38°9'S, 
61°48'W,  15.x.  1973  (Maury  &  Cesari,  MACN);  Sierra 
de  la  Ventana,  38°9'S,  61°48'W,  15.vii.l972  (Amarril- 
la,  MACN).  Cordoba:  C.  Paz,  31°24'S,  64°31'W, 
15.V.1940  (C.  Marti,  MACN);  Calamuchita,  32°4'S, 
64°33'W,  15.xii,1941  (J.  M.  Viana,  MACN),  15.i.l955 
(J.  M.  Viana,  MACN);  Mina  Clareo,  31°43'S,  65°0'W, 
15.iv.l973  (Stiebel,  MACN).  Jtijuij:  Iturbe,  22°59'S, 
65°21'W,  22.ii.1983  (L.  E.  Peila,  AMNH).  La  Rioja: 
Ilias  [?],  29°5'S,  66°19'W  (P.  M.  Gomez,  MACN). 
San  Luis:  Chosmes  and  Desaguadero  (Mendoza), 
33°24'30"S,  67°0'0"W,  14.iv.l967  (L.  E.  Pena,  MCZ). 
Santiago  del  Estero:  Santiago  del  Estero,  27°47'S, 
64°16'W  (AMNH),  3.iv.l965'"(H.  W.  Levi,  MCZ);  70 
km  W  Santiago,  27°47'S,  65°25'W,  3.iv.l965  (H.  W. 
Levi,  MCZ);  Quebrachos:  Sumampa,  Parayacu  [?], 
29°22'S,  63°28'W,  15.xi.l944  (Maldonado  Bruzzone, 
MLP). 

14.  Metepeira  galatheae  (Thorell) 
Figures  108-120,  304;  Map  7 

Epeira  galatheae  Thorell,  1891:  53.  Female  holotype 
from  "Cobija,  Bolivia,"  now,  Cobija,  Antofagasta, 
Chile,  in  the  UZMK,  examined. 

Araneus  galatheae: — Bonnet,  1955:  506. 

Metepeira  galatheae: — Levi,  1991:  179.  Platnick, 
1993. 

Metepeira  cereicola  nomen  nudum,  female  in  AMNH 
but  no  description  has  been  found.  Female  man- 
uscript type  from  Salamanca,  Coquimbo,  Chile, 
collected  by  Archer  on  30. iv.  1961. 

Note.  Thorell  (1891)  listed  the  holotype  s  locality 
as  "Cobija,  Bolivia."  While  Cobija,  Bolivia  exists 
(11°02'S"68°44'W),  it  is  an  unlikely  locality  for  the 
corvette  Galathea  to  visit  on  its  voyage  to  southern 
Asia  because  it  would  have  meant  climbing  over 
the  Andes.  Instead,  the  port  town  of  Cobija,  Chile 
(22°33'S  70°16'W)  is  much  more  likely,  especially 
since  this  region  of  Chile  was  under  Bolivian  ad- 
ministration throughout  the  period  of  Galathea's 
voyage,  1845-1847  (Paynter  et  al,  1975). 

Description.  Female  from  Chile  Chico, 
Aisen  province,  Chile.  Reddish  brown  car- 
apace with  white  setae,  light  around  eyes, 
lateral  posterior  extensions,  central  arrow- 
shaped  inark  (Fig.  113).  Legs  light  yellow, 
articles  annulated  distally.  Femur  I  with 
row  of  four  or  five  macrosetae  on  anterior 
side;  two  to  four  on  anteroventral  side. 
Dorsum  with  black  and  white  setae.  Foli- 
um speckled  browii  with  a  white  fleur-de- 
lis  that  reduces  posteriorly  (Fig.  113).  Ven- 
ter brownish  gray  with  wide  median  white 


line  and  pair  of  large  white  spots  on  either 
side  of  spiracle.  A  pair  of  very  faint  white 
stitching  parallel  to  and  on  either  side  of 
median  line  (Fig.  114).  Sternum  reddish 
brown,  sometimes  with  small  posterior  and 
anterior  white  marks  (Fig.  114).  Ratio  of 
eye  diameters:  posterior  medians  and  an- 
terior iTiedians  1.0,  anterior  laterals  1.2, 
posterior  laterals  1.2.  Anterior  median 
eyes  separated  by  1.7  diameters,  posterior 
median  eyes  by  1.2,  anterior  median  eyes 
separated  from  anterior  laterals  by  3.6  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.1  their  diameters.  To- 
tal length  7.8  mm.  Carapace  3.5  mm  long, 
2.7  wide.  First  femur  3.7  mm,  patella  and 
tibia  3.9,  metatarsus  3.5,  tarsus  1.2.  Sec- 
ond patella  and  tibia  3.4  mm,  third  2.1, 
fourth  3. 

Male  from  Chile  Chico,  Aisen  province, 
Chile.  Leg  coloration  as  in  feinale,  except 
femur  I  mostly  reddish  browii  with  row  of 
four  macrosetae  on  anterior  side;  five  on 
anteroventral  side.  Carapace,  dorsum,  ven- 
ter and  sternum  as  in  female  (Figs.  Ill, 
112).  Ratio  of  eye  diameters:  posterior  me- 
dians and  anterior  medians  1.0,  anterior 
laterals  1.3,  posterior  laterals  1.3.  Anterior 
median  eyes  separated  by  1.6  diameters, 
posterior  median  eyes  by  1.2,  anterior  me- 
dian eyes  separated  froixi  anterior  laterals 
by  3.3  diameters  of  anterior  lateral  eyes, 
lateral  eyes  separated  by  0.4  their  diame- 
ters. Total  length  5.3  mm.  Carapace  2.7 
mm  long,  2.1  wide.  First  femur  3.9  mixi, 
patella  and  tibia  3.8,  metatarsus  3.6,  tarsus 
1.1.  Second  patella  and  tibia  3.2  mm,  third 
1.8,  fourth  2.6. 

Diagnosis.  Although  the  sternum  of  M. 
galatheae  varies  from  solid  brown  (Fig. 
114)  to  brown  with  median  white  marks, 
the  venter  of  the  abdomen  is  surprisingly 
consistent.  The  venter  has  a  wide  brownish 
gray  area  and  a  short  median  white  line 
flanked  by  very  faint  indications  of  parallel 
lines  (Fig.  114).  In  contrast,  the  flanking 
lines  in  M.  karkii  thicken  anteriorly  (Fig. 
128)  and  the  venter  of  M.  nigriventris  is 
almost  completely  black  (Fig.  91).  Despite 
enormous   variation    in   the   hood   of  the 


44         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


-40  -35 

Latitude  (Degrees  North) 

Figure  109.  Days  on  which  mature  M.  galatheae  were  col- 
lected between  1937  and  1989  with  latitude  of  the  collection 
locality.  Seasonality  appears  to  be  more  restricted  in  southern 
regions  than  in  northern  regions. 

Scale  of  abscissa:  -150  =  August  3;  1  =  January  1;  150  = 
May  30. 


scape  with  a  corresponding  variation  in  the 
position  of  the  epigynal  openings,  it  is 
nonetheless  possible  to  distinguish  the  fe- 
males of  M.  galatheae  from  those  of  other 
species  in  the  M.  nigriventris  group. 
Whether  ventrally  or  posteriorly  posi- 
tioned, the  epigynal  openings  and  the 
darkened  shadows  of  the  sclerotized  re- 
ceptacles beneath  them  are  relatively 
smaller  than  those  of  M.  calamuchita,  M. 
tarapaca,  and  M.  nigriventris  (compare 
Figs.  118-120  with  Figs.  86,  94,  102).  Also, 
M.  galatheae  lacks  the  swollen  posterior 
lobes  present  in  M.  karkii  (compare  Figs. 
118-120  with  Fig.  123).  The  shape  of  the 
embolus  of  M.  galatheae  varies  significant- 
ly (compare  Fig.  109  with  Fig.  110).  How- 
ever, unlike  other  species,  the  embolus  of 
M.  galatheae  has  a  distinct,  round  and 
swollen  protrusion  (Fig.  117). 

Variation.  Average  body  length  of  80  fe- 
males examined  7.9  mm,  range  4.8  to  12.5 
mm.  Average  body  length  of  16  males  ex- 
amined 5.9  mm,  range  3.5  to  8.1  mm.  Epi- 
gyna  vary  considerably.  Many,  similar  to 
the  holotype,  open  ventrally  and  resemble 
a  posteriorly  widened  version  of  M.  tara- 
paca (Figs.  119,  120);  cleared  epigyna 
show  relatively  straight  ducts  connecting 
the  epigynal  openings  with  the  seminal  re- 
ceptacles. These  have  a  short  distance  be- 
tween the  openings  and  the  hood  of  the 
scape.  Others,  usually  in  southern  Chile, 
have  posterior  openings  and  look  surpris- 


ingly different  (Fig.  118);  cleared  epigyna 
show  S -shaped  ducts  connecting  the  open- 
ings with  the  seminal  receptacles.  These 
have  an  extended  wrinkled  area  between 
the  openings  and  the  hood  of  the  scape. 
However,  several  females  [e.g.,  CHILE 
Bio-Bio:  Chilian,  8.xi.l976  (G.  Moreno, 
AMNH);  Las  Lajuelas,  ll.i.l976  (G.  Mo- 
reno, AMNH)]  have  epigyna  that  appear 
to  be  intermediate  between  the  two  forms. 
Furthermore,  no  somatic  features  were 
found  to  be  sufficiently  different,  and  little 
corresponding  variation  was  found  among 
sympatric  males.  It  is  possible  that  further 
collecting  efforts  will  discover  correspond- 
ing males,  and  future  molecular  studies 
may  show  that  speciation  has,  in  fact,  oc- 
curred. But  in  the  meantime,  I  am  opting 
to  treat  both  varieties  as  belonging  to  the 
same  species. 

Natural  History.  Although  mature  spec- 
imens have  been  collected  throughout  the 
year  (Fig.  304),  the  seasonality  of  this  spe- 
cies appears  to  depend  on  its  latitude.  At 
the  45th  southern  parallel,  spiders  are  usu- 
ally found  in  late  November  and  Decem- 
ber; at  the  40th  parallel,  spiders  occur  be- 
tween October  and  Febrtiary;  and  at  the 
30th  parallel,  they  are  collected  year  round 
(Fig.  108).  Median  elevation,  550  m.  Spi- 
ders are  found  on  Patagonian  scrub, 
dunes,  and  wire  fences. 

Distribution.  Chile  and  Argentina  (Map 
7). 

Records  Examined.  ARGENTINA  Buenos  Aires: 
Felipe  Sola,  38°1'S,  62°50'W,  15.1.1944  (Prosen, 
MLP);  Patagones,  40°48'S,  62°59'W,  15.ii.l937  (J.  M. 
Viana,  MACN);  Sierra  de  la  Ventana,  38°9'S, 
61°48'W,  15.iii.l939  (J.  C.  Gario,  MACN).  Catamar- 
ca:  Mutquin,  28°19'S,  66°10'W,  15.1.1963  (O.  de  Fer- 
rariis,  AMNH).  Chubut:  15  km  S  Epuyen,  42°22'S, 
71°21'W,  15.1.1986  (P  A.  Goloboff,  N.  I.  Platnlck,  & 
R.  T.  Schuh,  AMNH);  19.5  km  E  Shaman,  44°27'S, 
70°30'W,  19.xi.l966  (E.  I.  Schllnger  &  M.  E.  Irwin, 
CAS);  3  km  N  Puerto  Lobos"  41°59'S,  65°6'W, 
14.xii.l966  (E.  I.  Schllnger  &  M.  E.  Irwin,  CAS);  35 
km  E  Esquel,  42°54'S,  70°53'W,  18.xl.l966  (E.  I. 
Schhnger  &  M.  E.  Irwin,  CAS);  El  Hoyo  [?],  42°4'S, 
71°30'W  (A.  Kovacs,  AMNH),  10.1.1962  (Andor  Ko- 
vacs,  AMNH);  Epuyen,  42°15'S,  71°23'W,  18.xl.l962 
(Andor  Kovacs,  AMNH);  Leleque,  42°28'S,  71°6'W, 
12.11.1965  (Andor  Kovacs,  AMNH);  Los  Manantlales, 


Metepeira  •  Piel 


45 


A    A 


Figures  110-120.  Metepeira  galatheae  {ThoreW)  (sp.  14  [110-114,116,117,120]  46°33'S,  71°57'W;  [115,119]  29°50'S,  70°2'W; 
[118]  33°30'S,  71°25'W).  110,  male  palpus,  mesal.  Ill,  male,  dorsal.  112,  male,  ventral.  113,  female,  dorsal.  114,  female, 
ventral.  115,  epigynum,  posterior.  116,  epigynum,  posterior.  117,  male  embolic  division,  ventral.  118-120,  epigynum,  ventral. 
Figures  121-128.  Metepeira  /car/f/V  (Tullgren)  (sp.  15;  51°38'S,  69°13'W).  121,  male  palpus,  mesal.  122,  epigynum,  posterior. 
123,  epigynum,  ventral.  124,  male,  dorsal.  125,  male,  ventral.  126,  female,  dorsal.  127,  female,  ventral. 
Scale  bars:  dorsum  and  venter  figures  1.0  mm. 


N  of  Comodoro-Rivadavia,  45°28'S,  69°29'W, 
19.xi.l985  (L.  E.  Peiia,  AMNH);  N  of  Camarones, 
Cantera,  Namuncura,  44°46'S,  65°42'W,  17..>d.l9S5 
(L.  E.  Pena,  AMNH);  Rio  Turbio,  42°13'S,  71°41'W 
(Andor  Kovacs,  AMNH),  12.1.1962  (Andor  Kovacs, 
AMNH).  Cordoba:  12  mi  W  Sampacho,  33°23'S, 
64°43'W,  7.ii.l951  (E.  S.  Ross  &  Michelbacher,  CAS); 
Arguello,  31°21'S,  64°15'W,  15.xii.l943  (J.  A.  De  Car- 


lo, MACN);  Calamuchita,  32°4'S,  64°33'W, 
15.xii.l940  (J.  M.  Viana,  MACN);  Sampacho,  33°23'S, 
64°43'W,  7.ii.l951  (E.  S.  Ross  &  Michelbacher,  CAS). 
Mendoza:  Between  Beazley  and  San  Rafael,  34°10'S, 
67°29'W,  4.iii.l983  (L.  E.  Pefia,  AMNH);  Mendoza, 
32°53'S,  68°49'W,  30.iii.l965  (H.  W.  Levi,  MCZ);  Us- 
pallata,  32°35'S,  69°20'W,  7.iii.l983  (L.  E.  Pefia, 
AMNH).  Neuquen:  Catan  Lil,  Charaliuilla,  39°45'S, 


46         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


70°37'W,  15.ii.l971  (O.  de  Ferrariis,  AMNH);  Cuba 
del  Leon  [?],  39°9'S,  70°53'W,  15.1.1975  (Mauiy, 
MACN);  Lago  Alumine,  38°55'S,  71°9'W,  15.1.1976 
(O.  de  Ferrariis,  AMNH);  Zapala,  38°54'S,  70°4'W, 
15.1.1958  (J.  R.  Navas,  MACN).  Rio  Negro:  Cerro 
Alto  [?],  41°8'S,  70°40'W  (MACN);  Co.  Leones, 
source  of  Rio  Limay,  40°33'S,  70°26'W,  28.11.1959  (J. 
R.  Navas,  MACN);  El  Bolson,  41°58'S,  71°31'W, 
1.11.1961  (A.  Kovacs,  AMNH),  17.X.1961  (Andor  Ko- 
vacs,  AMNH);  El  Bolson,  41°58'S,  71°35'W,  2.11.1965 
(Andor  Kovacs,  AMNH);  Ceneral  Roca,  39°3'S, 
67°32'W,  15.lx.1964  (Bachmann,  MEG).  Salta:  Mauiy 
[?],  24°40'S,  65°45'W,  15.1.1975  (MACN).  San  Juan: 
10  km  N  Matagusanos,  31°10'S,  68°38'W,  13.1.1983 
(L.  E.  Peiia,  AMNH).  San  Luis:  INTA  Experimental 
Station,  E  of  Villa  Mercedes,  33°40'S,  65°27'W, 
8.xil.l967  (C.  R.  Ward,  CAS).  Santa  Cruz:  2.4  km  S 
Fltz  Roy,  47°2'S,  67°15'W,  12.xll.l966  (E.  I.  Schllnger 
&  M.  E.  Irwin,  CAS).  Tucumdn:  San  Miguel  de  Tu- 
cuman,  IML  gardens,  26°49'S,  65°13'W,  19.xil.l979 
(L.  A.  Stange,  FSCA).  BRAZIL  Mato  Grosso:  Campo 
Grande,  20°27'S,  54°37'W,  7.11.1952  (M.  Alvarenga, 
MZSP).  CHILE  Aconcagua:  W  end  tunnel,  85  km  S 
Illapel,  32°49'S,  71°7'W,  29.xl.1950  (E.  S.  Ross  & 
Mlchelbacher,  CAS).  Alsen:  8  km  W  Chile  Chlco, 
46°33'S,  71°57'W,  22.xi.1966  (E.  I.  Schllnger  &  M. 
E.  Ii^wln,  CAS);  Chile  Chlco,  near  lake,  46°33'S, 
71°43'W,  21.xl.1966  (E.  I.  Schllnger  &  M.  E.  Invln, 
CAS).  Antofagasta:  6  km  N  Muelle  de  Pledra,  N  Tal- 
tal,  25°21'S,  70°30'W,  4.11.1942  (Junius  Bird,  AMNH); 
Caleta  Hueso  Parado,  Taltal,  25°22'S,  70°28'W, 
1.11.1941  (Junius  Bird,  AMNH);  Coblja,  22°33'S, 
70°16'W  (ZMUC);  Quebrada  Paposo,  25°2'S, 
70°27'W,  3.11.1989  (L.  Stange,  FSCA).  Araucania:  Pe- 
mehue  [?],  38°3'S,  71°43'W  (L.  E.  Pena,  IRSNB); 
Villarrlca,  36°16'S,  72°13'W,  25.xl.1963  (G.  F  Ed- 
munds, AMNH).  Atacama:  50-60  km  S  Copiapo, 
27°51'S,  70°20'W,  24.vlll.1966  (E.  I.  Schllnger  &  M. 
E.  Irwin,  CAS);  Copiapo,  27°22'S,  70°20'W  (Cartis, 
MNRJ);  Rio  Copiapo,  by  the  sea,  27°19'S,  70°56'W, 
13.vl.1968  (L.  E.  Pena,  MCZ).  Bio-Bio:  4  km  E  road 
to  Pinto,  36°42'S,  71°53'W,  4.1.1976  (B.  Moreno, 
AMNH);  Chilian,  36°36'S,  72°7'W,  2.1.1976  (G.  Mo- 
reno, AMNH),  21.11.1978  (G.  Moren,  MCZ);  Chilian, 
In  cemetery,  36°36'S,  72°7'W,  8.xl.l976  (G.  Moreno, 
AMNH);  Cuesta  de  QuUmo,  Chilian,  36°38'S, 
72°12'W,  13.xl.1976  (G.  Moreno,  AMNH);  El  Aban- 
Ico,  37°20'S,  71°31'W,  30.xll.l950  (E.  S.  Ross  & 
Mlchelbacher,  CAS);  Las  Lajuelas,  36°39'S,  72°8'W, 
11.1.1976  (G.  Moreno,  AMNH).  Concepclon, 
36°50'S,  73°3'W  (L.  E.  Pena,  IRSNB);  Concepclon: 
Salta  de  Rio  Laja,  37°13'S,  72°23'W,  30.1.1951  (E.  S. 
Ross  &  Mlchelbacher,  CAS);  Nuble:  50  km  E  San 
Carlos,  36°25'S,  71°6'W,  26.xll.1950  (E.  S.  Ross  & 
Mlchelbacher,  CAS);  Nuble:  Cordillera  de  Chilian 
[?],  36°51'S,  7r24'W,  1.11.1947  (L.  E.  Peiia,  IRSNB); 
Rio  Andallen,  36°44'S,  73°1'W,  25.111.1979  (S.  Gu- 
tierrez, MCZ).  Coquinibo:  20  ml  E  La  Serena, 
29°54'S,  70°56'W,  3.vll.l950  (E.  S.  Ross  &  Mlchel- 
bacher, CAS);  5  ml  N  Ovalle,  30°31'S,  71°12'W, 
l.xil.l950  (E.  S.  Ross  &  Mlchelbacher,  CAS);  Banos 


del  Toro,  29°50'S,  70°2'W,  15.11.1947  (L.  E.  Pena, 
IRSNB);  Cerro  Tahnay,  30°50'29"S,  71°37'14"W, 
29.xl.1961  (A.  F.  Archer,  AMNH);  Cuesta  las  Cardas, 
Ovalle  Rd.,  30°17'S,  71°16'W,  13.xl.l961  (R.  Wagen- 
knecht,  AMNH);  Hacienda  Illapel,  31°36'S,  71°7'W, 
3.X1.1954  (L.  E.  Pena,  IRSNB),  19..X.1966  (E.  I. 
SchUnger,  M.  E.  Irwin,  &  L.  E.  Pena,  CAS);  Illapel: 
Salamanca:  Fundo  Quelen,  31°52'S,  70°52'W, 
30.lv.1961  (A.  F.  Archer,  AMNH);  La  Serena, 
29°54'28"S,  7ri5'15"W,  15.11.1947  (L.  E.  Pena, 
IRSNB);  Loma  de  Penuelas,  6  km  S  La  Serena, 
29°57'S,  71°18'W,  28.xl.1961  (A.  F.  Archer,  AMNH); 
Qullacan,  16  km  E  La  Serena,  29°54'S,  71°5'W, 
2.x.  1961  (R.  Wagenknecht,  AMNH).  Los  Lagos:  Purr- 
anque,  40°55'S,  73°10'W,  15.11.1955  (Edwin  Reed, 
AMNH);  Rio  Bueno,  40°19'S,  72°58'W  (L.  E.  Pena, 
IRSNB);  Valdlvla:  Neltume,  39°48'S,  71°57'W, 
23.xi.1988  (V.  &  B.  Roth,  CAS).  Malleco:  Angol, 
37°48'S,  72°43'W,  29.1.1951  (E.  S.  Ross  &  Mlchel- 
bacher, CAS).  Maule:  Linares,  35°51'S,  71°36'W  (L. 
E.  Pena,  IRSNB),  15.1.1947  (L.  E.  Pena,  IRSNB); 
Mlraflores,  Pedag.  [?],  35°55'S,  71°39'W  (Toro, 
AMNH).  O'Higgins:  Cheplca,  34°44'S,  71°17'W, 
15..X11.1947  (L.  E.  Peiia,  IRSNB).  Region  Metropoli- 
tana:  34  km  W  Sandago,  33°30'S,  71°25'W, 
19.xll.1950  (E.  S.  Ross  &  Mlchelbacher,  CAS);  Ba- 
tuco,  nr.  Santiago,  33°13'S,  70°47'W  (Gull.  Mann, 
AMNH);  Lampa,  33°17'S,  70°54'W,  l.v.1979  (L.  E. 
Peiia,  AMNH).  Santiago:  El  Golf  [?],  33°30'S, 
71°25'W,  9.iv.l961  (A.  F  Archer  &  J.  Aros,  AMNH); 
Santiago,  33°30'S,  71°25'W  (L.  E.  Peiia,  IRSNB), 
1.11.1973  (W.  C.  Sedgvdck,  MCZ).  Talca:  22  ml  N  Tal- 
ca,  35°7'S,  71°40'W,  22.xll.1950  (CAS).  Valparaiso: 
Concon,  in  cow  farm,  32°55'S,  71°31'W,  4.111.1962  (H. 
Morales,  AMNH);  La  Cruz,  32°53'S,  71°16'W, 
18.1.1973  (W.  C.  Sedgwick,  MCZ);  Llay-Llay,  32°51'S, 
70°58'W,  20.1.1973  (W.  C.  Sedgwick,  MCZ);  Los  Mai- 
tenes  [?],  32°59'S,  71°15'W,  14.X.1954  (L.  E.  Peiiai 
IRSNB);  Quintay,  33°11'S,  71°42'W,  19.11.1967  (E.  I. 
Schhnger,  CAS);  Valparaiso,  33°2'S,  71°38'W  (Edwin 
Reed,  AMNH). 

15.  Metepeira  /car/c/7  (Tullgren) 
Figures  121-128,  303;  Map  5 

Araneus  karkii  Tullgren,  1901:  219,  259.  Female  lio- 
lotype  from  Kark,  Chile  in  the  SMNH,  examined. 

Metepeira  labyrinthea: — Roewer,  1942:  868.  Bonnet, 
1957:  2821.  Erroneous  synonymy. 

Description.  Female  from  Rio  Gallegos, 
Santa  Cruz  Province,  Argentina.  Carapace 
reddish  brown  with  white  setae,  light 
around  eyes  with  lateral  posterior  exten- 
sions (Fig.  127).  Legs  light  yellow,  articles 
annulated  distally.  Femur  I  with  row  of 
three  to  four  macrosetae  on  anterior  side; 
one  to  four  on  anteroventral  side.  Dorsum 
covered   with    denser,    longer,    black   and 


Metepeira  •  Piel 


47 


white  setae  than  in  most  species.  Folium 
mostly  white  with  brown  speckles  (Fig. 
127).  Venter  brownish  gray  with  wide 
white  median  line;  pair  of  large  white  spots 
on  either  side  of  spiracle.  A  pair  of  thin 
white  lines,  parallel  to  and  on  either  side 
of  median  line,  sometimes  connect  to  a 
pair  of  thinner  transverse  white  lines:  one 
just  posterior  to  the  epigynal  groove,  one 
just  anterior  to  the  spinnerets  (Fig.  128). 
Sternum  dark  reddish  brown  (Fig.  128). 
Ratio  of  eye  diameters:  posterior  inedians 
and  anterior  medians  1.2,  anterior  laterals 
1.4,  posterior  laterals  1.5.  Anterior  median 
eyes  separated  by  1.8  diameters,  posterior 
inedian  eyes  by  1.0,  anterior  median  eyes 
separated  from  anterior  laterals  by  4.2  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.3  their  diameters.  To- 
tal length  9.5  inm.  Carapace  4  mm  long, 
3.4  wide.  First  femur  4  mm,  patella  and 
tibia  4.4,  metatarsus  3.7,  tarsus  1.2.  Sec- 
ond patella  and  tibia  3.9  mm,  third  2.5, 
fourth  3.8. 

Male  from  Rio  Gallegos,  Santa  Cruz 
Province,  Argentina.  Carapace  reddish 
brown  with  lighter  eye  region,  lateral  pos- 
terior extensions,  and  median  arrowhead 
inark  (Fig.  125).  Legs  light  yellow,  articles 
distally  annulated  reddish  brown.  Femur  I 
with  row  of  three  to  four  macrosetae  on 
anterior  side;  five  to  six  on  anteroventral 
side.  Dorsal  folium  mostly  white  with 
brown  speckles  (Fig.  125).  Venter  and 
sternuiTi  as  in  female  (Fig.  126).  Ratio  of 
eye  diameters:  posterior  medians  and  an- 
terior medians  1.0,  anterior  laterals  1.3, 
posterior  laterals  1.2.  Anterior  inedian 
eyes  separated  by  1.7  diameters,  posterior 
inedian  eyes  by  1.2,  anterior  median  eyes 
separated  from  anterior  laterals  by  2.8  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.4  their  diameters.  To- 
tal length  5.7  mm.  Carapace  2.8  mm  long, 
2  wide.  First  femur  4.3  mm,  patella  and 
tibia  4.3,  metatarsus  4.1,  tarsus  1.2.  Sec- 
ond patella  and  tibia  3.5  mm,  third  1.9, 
fourth  2.9. 

Diagnosis.  Females  are  easily  separated 
from  other  species  in  the  M.  nigriventris 


group  by  the  thick  posterior  epigynal  lobes 
(compare  Fig.  123  with  Fig.  119).  The  dis- 
tal embolic  apophysis  does  not  protrude 
out  from  under  the  terminal  apophysis 
(Fig.  124)  as  it  does  in  M.  galatheae,  M. 
tarapaca,  and  M.  nigriventris  (Figs.  87,  95, 
119).  The  embolus  of  M.  karkii  differs 
from  M.  calaniuchita  by  lacking  the  in- 
wardly curved  "dewlap"  under  the  embo- 
lus (compare  Fig.  121  with  Fig.  103). 

Variation.  Average  body  length  of  13  fe- 
males examined  6.8  mm,  range  5  to  8.2 
mm.  Average  body  length  of  five  males  ex- 
amined 4.4  mm,  range  2  to  5.3  mm.  Dor- 
sal folia  vaiy  from  white  with  little  contrast 
and  indistinct  fleur-de-lis  to  darker  with 
more  contrast  and  distinct  fleur-de-lis. 

Natural  History.  This  species  appears  to 
be  strongly  seasonal:  mature  specimens 
have  been  collected  between  November 
and  March  (Fig.  303).  Median  elevation, 
300  m.  Spiders  are  found  in  pampas  (tree- 
less grassland). 

Distribution.  Lower  altitudes  in  south- 
ern Argentina  and  Chile  (Map  5). 

Records  Examined.  ARGENTINA  Chubut:  Puerto 
Piramides,  Peninsula  Valdes,  42°34'S,  64°17'W, 
12.xi.l988  (V.  &  B.  Rodi,  CAS).  Neuqtien:  Laguna 
Blanca,  39°3'S,  70°23'W,  15.iii.l959  (J.  Nara, 
MACN);  Zapala,  38°54'S,  70°4'W,  15.1.1958  (J.  R.  Na- 
vas,  MACN);  Zapala,  Laguna  Blanca,  38°54'S, 
70°4'W,  15.1.1959  (J.  R.  Navas,  MACN).  Rto  Negro: 
Cerro  Alto  [?],  41°8'S,  70°40'W  (MACN);  Coronel 
Juan  Jose  Comez,  39°2'S,  67°39'W,  15..\i.l945  (Ibarra 
Grasso,  MLP);  Ne-Luan,  41°25'S,  68°45'W  (MACN). 
Santa  Cruz:  Laguna  Calafate,  Precordllfera  [?], 
50°55'S,  70°9'W,  22.1.1967  (P  San  Martin,  MCZ);  Rio 
Gallegos,  51°38'S,  69°13'W,  20.1.1967  (P  San  Martin, 
MCZ).  BOLIVIA  Santa  Cniz:  Patagonia:  Estancia 
Monte,  cerca  Rio  Coyby  [?],  50°14'S,  68°55'W  (B. 
Brown,  AMNH).  CHILE  Magallanes:  4  km  W  La- 
guna Amarga,  50°59'S,  72°49'W,  8.xil.l966  (E.  I. 
Schlinger  &  M.  E.  Irwin,  CAS);  Kark,  51°17'30"S, 
72°32'30"W,  13.111.1899  (E.  Nordenskiold,  NRMS). 

Metepeira  compsa  Group 

The  three  species  in  the  M.  co77ipsa 
group  include  Metepeira  compsa,  Metepei- 
ra roraima,  and  Metepeira  gressa.  Along 
with  the  Metepeira  nigriventris  group,  this 
group  has  a  median  apophysis  with  teeth 
on  the  face  of  the  keel.  Unlike  the  Mete- 


48         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


peira  nigriventris  group,  this  group  has 
smaller,  slimmer  distal  embolic  apophyses 
that  do  not  arch  up  over  the  embolus  tip 
(compare  Fig.  149  with  Fig.  84).  The 
openings  to  the  epigynum  have  distinct, 
sclerotized  edges  and  are  clearly  visible  ei- 
ther as  circles  (Figs.  131,  134)  or  eye- 
shaped  ovals  (Figs.  143,  151). 

1 6.  Metepeira  compsa  (Chamberlin) 
Figures  129-140,  311;  Map  8 

Aranea  compsa  Chamberlin,  1916:  252,  fig.  6,  ? .  Fe- 
male holotype  from  Ollantaitambo,  Cusco,  Peru,  in 
the  MCZ,  examined.  Bonnet,  1955:  462. 

Araneus  lahijrintheus: — Petnmkevitch,  1926:  27.  Er- 
roneous synonymy. 

Metepeira  virginensis  Chamberlin  and  Ivie,  1942:  70, 
figs.  188-190,  9,8.  Female  holotype  from  St. 
Thomas,  U.S.  Virgin  Islands,  in  AMNH,  examined. 
NEW  SYNONYMY. 

Metepeira  latigijna  Chamberlin  and  Ivie,  1942:  70, 
figs.  191,  192,  5  .  Female  holotype  from  Porto  Ale- 
gre  [Baliia],  Brazil,  in  AMNH,  examined.  NEW 
SYNONYMY. 

Metepeira  compsa: — Chamberlin  and  Ivie,  1942:  71, 
figs.  193-195,  $,  c?. 

Metepeira  labyrinthea: — Biyant,  1942:  346. 

Metepeira  perezi  Archer,  1965:  132,  figs.  14,  16,  3, 
9 .  Male  holotype  from  subexperiment  station,  Is- 
abela,  Puerto  Rico,  in  AMNH,  examined.  Brignoli, 
1983:  276.  NEW  SYNONYMY. 

Metepeira  vaurieonim  Archer,  1965:  133,  figs.  15,  19, 
S,  ?.  Male  holotype  from  Usine  de  Robert,  Mar- 
tinique, in  AMNH,  lost.  BrignoU,  1983:  276.  Lo- 
pez, 1993:  10,  11,  figs.  1-4,  11,  6.  NEW  SYN- 
ONYMY. 

Note.  Although  the  type  for  M.  vaurieonim  is 
lost,  all  examined  records  from  Martinique  (includ- 
ing ones  from  "Usine  de  Robert")  belong  to  M. 
compsa. 

Description.  Female  from  Savonet,  Cu- 
rasao, Netherlands  Antilles.  Carapace  light 
around  eyes  with  lateral  posterior  exten- 
sions (Figs.  135,  137).  Only  tibia  IV 
ringed.  Femur  I  with  row  of  four  macro- 
setae  on  anterior  side;  two  or  three  fine 
setae  on  anteroventral  side.  Dorsum  of  ab- 
domen with  usual  Metepeira  folium, 
though  whiter  than  usual  in  some  speci- 
mens (Figs.  135,  137).  Venter  with  wide 
white  median  line  flanked  by  two  thin 
(Fig.  136)  or  wide  (Fig.  138)  white  lines; 
pair  of  white  spots  on  either  side  of  spi- 
racle. Sternum  has  wide  median  white  line 


widening  anteriorly,  sometimes  broken 
(Figs.  136,  138).  Ratio  of  eye  diameters: 
posterior  medians  and  anterior  medians 
1.0,  anterior  laterals  1.3,  posterior  laterals 
1.2.  Anterior  median  eyes  separated  by  1.3 
diameters,  posterior  median  eyes  by  0.7, 
anterior  median  eyes  separated  from  an- 
terior laterals  by  1.8  diameters  of  anterior 
lateral  eyes,  lateral  eyes  separated  by  0.2 
their  diameters.  Total  length  3.9  mm.  Car- 
apace 1.8  mm  long,  1.4  wide.  First  femur 
1.9  mm,  patella  and  tibia  1.9,  metatarsus 
1.5,  tarsus  0.7.  Second  patella  and  tibia  1.7 
inm,  third  0.9,  fourth  1.5. 

Male  from  Savonet,  Curasao,  Nether- 
lands Antilles.  Carapace  light  around  eyes 
with  lateral  posterior  extensions.  Slightly 
lighter  median  triangular  mark  anterior  to 
thoracic  furrow  (Fig.  139).  Legs  lightly 
ringed.  Femur  I  with  row  of  four  macro- 
setae  on  anterior  side,  four  on  anteroven- 
tral side.  Dorsum,  venter,  and  sternum  as 
in  female  (Figs.  139,  140).  Ratio  of  eye 
diameters:  posterior  medians  and  anterior 
medians  1.0,  anterior  laterals  1.3,  posterior 
laterals  1.2.  Anterior  median  eyes  separat- 
ed by  1.3  diameters,  posterior  median  eyes 
by  0.6,  anterior  median  eyes  separated 
from  anterior  laterals  by  1.4  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.3  their  diameters.  Total  length  3.6 
mm.  Carapace  1.8  min  long,  1.4  wide. 
First  femur  2.7  mm,  patella  and  tibia  2.8, 
metatarsus  2.4,  tarsus  0.9.  Second  patella 
and  tibia  2.3  mm,  third  1.2,  fourth  1.8. 

Diagnosis.  Unlike  other  species,  the 
openings  to  the  epigynum  of  M.  compsa 
are  small  and  almost  perfectly  round  and 
sclerotized  around  the  riin  (Fig.  131).  In 
Peruvian  and  Argentinean  populations  the 
scape  can  be  extremely  wide,  often  entire- 
ly covering  a  ventral  view  of  the  openings 
(Fig.  134).  The  distal  embolic  apophysis 
on  the  male  palp  extends  straight  from  its 
base  and  parallel  to  the  embolus  tip  (Figs. 
129,  132),  in  contrast  to  those  of  other  spe- 
cies in  the  M.  compsa  species  group  that 
lift  away  from  the  embolus  tip  (Figs.  141, 
149). 

Variation.  Average  body  length  of  22  fe- 


Metepeira  •  Piel        49 


roraima 
(17) 


Figures  129-140.  Metepeira  compsa  (Chamberlin)  (sp.  16  [129-131,135,136,139,140]  12°20'N,  69°7'W;  [132]  4°30'S,  81°8'W; 

[133,134]  4°51'S,  80°46'W;  [137,138]  17°5'N,  61°42'W).  129,  male  palpus,  mesal.  130,  epigynum,  posterior.  131,  epigynum, 

ventral.  132,  male  palpus,  mesal.  133,  epigynum,  posterior.  134,  epigynum,  ventral.  135,  female,  dorsal.  136,  female,  ventral. 

137,  female,  dorsal.  138,  female,  ventral.  139,  male,  dorsal.  140,  male,  ventral. 

Figures  141-148.  Metepeira  roraima  new  species  (sp.  17  [141-147]  3°22'N,  60°19'W;  [148]3°21'N,  76°33'W).  141,  male  palpus, 

mesal.  142,  epigynum,  posterior.  143,  epigynum,  ventral.  144,  male,  dorsal.  145,  male,  ventral.  146,  female,  dorsal.  147,  female, 

ventral.  148,  female,  dorsal. 

Scale  bars:  dorsum  and  venter  figures  1 .0  mm. 


males  examined  5  mm,  range  3.5  to  7.3 
mm.  Average  body  length  of  18  males  ex- 
amined 3.6  mm,  range  2.4  to  4.7  mm. 
Enormous  variation  in  the  shape  of  the 
scape  can  be  seen  in  this  species:  popula- 
tions in  the  Caribbean  and  northeastern 
South  America  have  a  narrow  scape  (Fig. 
131),    whereas    populations    in    Argentina 


and  Peru  have  a  thick  scape  (Fig.  134). 
Somewhat  less  consistently  parallel  differ- 
ences can  be  seen  in  the  shape  of  the  me- 
dian apophysis:  flagella  are  centered  in  Ca- 
ribbean and  northeastern  South  America 
(Fig.  129)  but  shifted  to  the  left  in  Argen- 
tina and  Peru  (Fig.  132).  Further  subtle 
differences  appear  in  the  shape  of  the  em- 


50         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


bolus  tip  and  embolus  arm  (compare  Fig. 
129  with  Fig.  132).  These  male  and  female 
genital  differences  are  not  sufficiently  con- 
sistent to  provide  solid  evidence  for  spe- 
ciation;  thus,  these  populations  will  be 
treated  as  one  species. 

Natural  History.  Mature  speciinens 
have  been  collected  throughout  the  year 
(Fig.  311).  Altitudes  range  from  just  above 
sea  level  (for  Caribbean  and  eastern  South 
American  localities)  to  4,000  m  (for  Pe- 
ruvian localities).  Spiders  have  been  found 
in  everything  from  second  growth  mixed 
exotics  (e.g.,  mango,  citrus,  banana)  and 
mangroves  to  grasses  and  semidesert  chap- 
arral. 

Distribution.  Puerto  Rico  to  northern 
Argentina,  but  absent  from  the  Amazon 
(Map  8). 

Records  Examined.  ARGENTINA  Buenos  Aires: 
Punta  Lara,  34°49'S,  57°59'W,  15.ii.l941  (F.  Moneos, 
MACN).  Chaco:  Basail,  27°52'S,  59°18'W,  18.iv.l942 
(MACN).  Corrientes:  Paso  de  la  Patria,  27°19'S, 
58°35'W,  15.1.1966  (M.  E.  Galiano,  MEG).  Entre 
Rws:  Parana,  31°44'S,  60°32'W  (Rosenzwaig,  MLP); 
Salto  Grande,  31°13'S,  57°56'W,  15.iii.l964  (M.  E. 
Galiano,  MEG).  Misiones:  Posadas,  27°23'S,  55°53'W, 
15.ix.l963  (M.  E.  Galiano,  MEG).  Neuquen:  Piedra 
del  Aguila,  40°3'S,  70°5'W,  18.vii.l978  (Mision  Cien- 
tifica  Danesa,  ZMUC).  Santa  Fe:  Tostado,  29°14'S, 
61°46'W  (A.  Giai,  MACN).  BOLIVIA  La  Paz:  Apolo, 
14°43'S,  68°31'W,  10.viii.l989  (L.  E.  Pefia,  AMNH). 
BRAZIL  Bahia:  Arquipelago  dos  Abrolhos,  17°40'S, 
38°50'W,  28.xii.1887  (U.S. E.G.,  Voy.  of  Albatross, 
USNM);  Parque  Ondina,  Salvador,  12°59'S,  38°31'W, 
25.vii.1962  (A.  F.  Archer,  AMNH);  Porto  Alegre, 
18°5'S,  39°34'W  (AMNH).  Minas  Gerais:  Pedra  /lzuI, 
16°1'S,  41°16'W,  15.xii.l970  (F.  M.  Oliveira,  AMNH). 
Periiambuco:  Pernambuco  [?],  8°3'S,  34°54'W, 
12.iii.l927  (SMF),  8.iii.l955  (SMF).  Rio  de  Janeiro: 
Lagomar  [?],  Macae,  near  sea,  22°23'S,  41°47'W, 
17.vii.l986  (R.  L.  C.  Baptista,  MZSP).  Rio  Grande  do 
Sal:  Montenegro,  29°42'S,  51°28'W,  3.xi.l977  (E.  H. 
Buckup,  MGN);  Rambo  [?],  30°4'0"S,  51°11'W 
(MNRJ);  Sao  Leopoldo,  29°46'S,  51°9'W,  14.X.1965 
(Gelia  Valle,  MZSP).  Sao  Paulo:  Rubiao,  Jr.  [?],  Bo- 
tucatu,  22°52'S,  48°26'W,  25.iv.1988  (R.  L.  G.  Bap- 
tista, MZSP);  Sao  Paulo,  Guamja,  24°0'S,  46°16'W, 
24.vii.1983  (R.  Sievers,  AMNH).  BRITISH  WEST 
INDIES  Anegada:  18°45'N,  64°20'W,  12.xi.l966 
(Harry  Beatty,  AMNH);  center,  nr.  salt  pond, 
18°45'N,  64°20'W,  4.vi.l966  (Island  Project  Staff, 
Univ  of  Puerto  Rico,  AMNH);  West  end,  18°45'N, 
64°22'W  4.vi.l966  (Island  Project  Staff,  Univ.  of 
Puerto  Rico,  AMNH);  Anegada  Settlement:  18°45'N, 
64°20'W,    5.vi.l966    (Island   Project    Staff,    Univ.    of 


Puerto  Rico,  AMNH).  Antigua:  Goolidge  Airport, 
17°6'N,  61°51'W,  15.xi.l967  (N.  L.  H.  Krauss, 
AMNH);  Devils  Bridge,  17°5'N,  61°42'W,  30.vi.l963 
(Rick  &  E.  N.  Kjellesvig-Waering,  AMNH);  Jolly 
Beach,  17°4'N,  61°53'W  20.ix.l963  (E.  N.  Kjellesvig- 
Waering,  AMNH);  Lignum  Vitae  Bay:  Jolly  Beach  [?], 
17°4'N,  61°53'W,  19.vi.l968  (E.  N.  Kjellesvig-Waer- 
ing, AMNH);  Redonda  Island,  from  webs  spun  be- 
tween boulders  on  beach,  16°55'N,  62°19'W, 
10.ivl9.56  (J.  F  G.  Glarke,  USNM);  Reeds  Point,  nr. 
Jolly  Beach;  17°4'N,  61°53'W,  2.vii.l963  (E.  N.  Kjel- 
lesvig-Waering, AMNH);  Saint  John's,  17°6'N, 
61°51'W,  15.viii.l967  (N.  L.  H.  Krauss,  AMNH), 
15.xii.l967  (N.  L.  H.  Krauss,  AMNH).  Dead  Mans 
Ghest  [?]:  18°22'N,  64°.34'W,  26.vl966  (Island  Pro- 
ject Staff,  Univ.  of  Puerto  Rico,  AMNH).  East  Seal 
Dog:  18°30'N,  64°25'W  7.vi.l966  (Island  Project 
Staff,  Univ.  of  Puerto  Rico,  AMNH).  George  Dog  Is- 
land: 18°30'N,  64°27'W,  7.vi.l966  (Island  Project 
Staff,  Univ.  of  Puerto  Rico,  AMNH);  4  mi  W  Virgin 
Gorda  Isl,  high  on  diy  cliffs,  18°30'N,  64°27'W, 
25.iii.1979  (K.  Johnson,  AMNH).  Ginger  Island: 
18°24'N,  64°28'W,  25.V.1966  (Island  Project  Staff, 
Univ  of  Puerto  Rico,  AMNH).  Great  Dog:  18°29'N, 
64°27'W,  7.vi.l966  (Island  Project  Staff,  Univ.  of 
Puerto  Rico,  AMNH).  Green  Cay,  near  Tortola: 
18°27'N,  64°42'30'W,  14.viii.l965  (Island  Project 
Staff,  Univ.  of  Puerto  Rico,  AMNH).  Island  O.  near 
Anegada  Setdement:  18°45'N,  64°20'W,  5.vi.l966  (Is- 
land'Project  Staff,  Univ.  of  Puerto  Rico,  AMNH).  Is- 
land R.  near  Anegada  Settlement:  18°45'N,  64°20'W, 
4.vi.l966  (Island  Project  Staff,  Univ.  of  Puerto  Rico, 
AMNH).  ]ost  Van  Dyke:  18°28'N,  64°45'W, 
30.viii.l965  (H.  Heatwole,  R.  Levins  &  F  Mac- 
Kenzie,  AMNH).  Large  mangrove  patch  nr.  Settle- 
ment Anegado:  18°45'N,  64°20'W  [?],  5.vi.l966  (Is- 
land Project  Staff,  Univ.  of  Puerto  Rico,  AMNH).  Lit-\ 
tie  Camanoe:  18°28'N,  64°33'W,  25.vl966  (Island 
Project  Staff,  Univ.  of  Puerto  Rico,  AMNH).  Little 
Josi  Van  Dyke:  18°27'N,  64°43'W,  27.vii.1965  (Island 
Project  Staff,  Univ  of  Puerto  Rico,  AMNH).  Little 
Tobago:  18°26'N,  64°51'W  4.iv.l966  (Island  Project 
Staff,  Univ.  of  Puerto  Rico,  AMNH).  Montserrat: 
Gage's  Soufriere  [?],  16°43'N,  62°10'W  28.vii.1972 
(N.  L.  H.  Krauss,  AMNH);  Plymouth,  16°42'N, 
62°13'W,  15.xi.l967  (N.  L.  H.  Krauss,  AMNH).  Neck- 
er  Island:  18°33'N,  64°21'W,  6.vi.l966  (Island  Project 
Staff,  Univ.  of  Puerto  Rico,  AMNH).  nr.  Anegada  Set- 
tlement: Byer's  Bache  [?],  18°45'N,  64°20'W, 
5.vi.l966  (Island  Project  Staff,  Univ.  of  Puerto  Rico, 
AMNH).  Peter  Island:  18°22'N,  64°35'W,  6.vii.l965 
(Island  Project  Staff,  Univ  of  Puerto  Rico,  AMNH), 
12.V.1966  (Percy  Chubb,  AMNH).  Prickly  Pear  Is- 
land: 17°10'N,  61°48'W,  6.vi.l966  (Island  Project 
Staff,  Univ.  of  Puerto  Rico,  AMNH).  Saint  Christo- 
pher Island:  Basseterre,  17°18'N,  62°43'W,  6.ii.l968 
(B.  Malkin,  AMNH).  Salt  Island:  18°23'N,  64°31'W, 
24.V.1966  (Island  Project  Staff,  Univ.  of  Puerto  Rico, 
AMNH).  Sandy  Key:  near  Tortola,  18°13'N,  63°7'W, 
31.viii.l965  (H.  &  A.  Heatwole,  AMNH).  Tobago  Is- 
land:   18°27'N,    64°48'W,    2.iv.l966    (Island   Project 


Metepeira  •  Piel 


51 


Staff,  Univ.  of  Puerto  Rico,  AMNH).  Tortola: 
18°27'N,  64°36'VV,  8.vii.l958  (A.  F.  Archer,  C.  Hel- 
sley,  &  M.  Sanderson,  AMNH);  Beef  Island,  18°27'N, 
64°31'W,  21.vii.l965  (H.  Heatwole,  R.  Levins  &  F. 
MacKenzie,  AMNH);  Greater  Camanoe  Isl,  18°29'N, 
64°32'W,  l.vii.l965  (Island  Project  Staff,  Univ.  of 
Puerto  Rico,  AMNH);  Little  Thatch  Island,  18°23'N, 
64°42'W,  16.viii.l965  (Island  Project  Staff,  Univ.  of 
Puerto  Rico,  AMNH);  Long  Bay  Estate,  18°24'N, 
64°41'W,  24.vii.1965  (Island  Project  Staff,  Univ.  of 
Puerto  Rico,  AMNH);  Marina  Key  [?],  18°27'N, 
64°36'W,  4.vii.l965  (H.  Heatwole  &  R.  Levins, 
AMNH);  Prospect  Reef,  S  Roadtown,  18°25'N, 
64°36'W,  23.iii.1979  (K.  Johnson,  AMNH),  31.iii.l979 
(K.  Johnson,  AMNH);  Road  to  town,  18°27'N, 
64°36'W,  15.vii.l972  (N.  L.  H.  Krauss,  AMNPI); 
Sandy  Spit  [?],  18°27'N,  64°36'W,  14.viii.l965  (Island 
Project  Staff,  Univ.  of  Puerto  Rico,  AMNH).  Toi-tola 
Island:  18°27'N,  64°36'W,  15.vii.l965  (Island  Project 
Staff,  Univ  of  Puerto  Rico,  AMNH).  Virgin  Gorda: 
18°30'N,  64°24'W,  15..xi.l966  (Hany  Beatty,  AMNH); 
Baths  &  Devils  Bay,  18°26'N,  64°27'W,  25.vi.1966 
(Island  Project  Staff,  Univ.  of  Puerto  Rico,  AMNH); 
Cooper  Mine  Trail  18°26'N,  64°26'W,  25.vi.1966  (Is- 
land Project  Staff,  Univ.  of  Puerto  Rico,  AMNH);  Sa- 
vana  Bay  &  Pond  Bay,  18°28'N,  64°25'W,  27.vi.1966 
(Island  Project  Staff,  Univ.  of  Puerto  Rico,  AMNH); 
Trelhs  Bay  (W  end  of  island),  nr.  harbor,  18°27'N, 
64°26'W,  27.iii.1979  (K.  Johnson,  AMNH);  Virgin 
Gorda  Mountain,  18°30'N,  64°24'W,  26.vi.1966  (Is- 
land Project  Staff,  Univ.  of  Puerto  Rico.  AMNH). 
Virgin  Islands  Area:  Island  Q  [?],  18°45'N,  64°20'W, 
5.vi.l966  (Island  Project  Staff,  Univ.  of  Puerto  Rico, 
AMNH).  West  Seal  'Dog  Island:  18°29'N,  64°28'W, 
7.vi.l966  (Island  Project  Staff,  Univ.  of  Puerto  Rico, 
AMNH).  CHILE  Tarapaca:  Arica,  18°29'S,  70°20'W, 
28.i.l973  (W.  C.  Sedgwick,  MCZ);  Azapa  Arica, 
18°31'S,  70°11'W,  9..xi.l955  (L.  E.  Peiia,  IRSNB); 
Lluta,  18°24'S,  70°19'W,  12.xi.l955  (L.  E.  Pena, 
IRSNB);  Rio  Lluta,  18°24'S,  70°19'W,  I2.xi.l955  (L. 
E.  Pena,  IRSNB);  Sobraya,  18°32'S,  70°9'W, 
10.xi.l955  (L.  E.  Peila,  IRSNB);  Taltape,  Camarones 
Valley,  18°59'S,  69°47'W,  29.i.l973  (MCZ).  COLOM- 
BIA Magdalena:  Cabaila  "Villa  Culebra,"  10  km  E 
Station  Marta,  11°12'N,  74°7'W,  15.X.1985  (H.-G. 
Miiller,  SMF);  Casajera  [?],  11°0'N,  74°15'W, 
l.ii.l974  (J.  A.  Kochalka,  JAK);  Cienaga,  11°1'N, 
74°15'W,  30.i.l974  (J.  A.  Kochalka,  JAK);  Gaira, 
11°11'N,  74°13'W,  15.xii.l975  (W.  Eberhard,  MCZ). 
DOMINICAN  REPUBLIC  Barahona:  Patos, 
10°38'N,  61°52'W,  24.ix.1944  (R.  H.  Montgomery, 
AMNH).  FRENCH  WEST  INDIES  Guadeloupe: 
Deshaies,  16°18'N,  61°48'W,  28.vi.1960  (C.  &  P  Vau- 
rie,  AMNH);  Domaine  Duclos  [?],  16°16'N,  61°31'W, 
25.vi.1960  (C.  &  P  Vaurie,  AMNH),  15.vii.l960  (C. 
&  P.  Vaurie,  AMNH);  Marie-Galante,  in  citrus, 
15°56'N,  61°16'W,  15.iii.l977  (W.  H.  Whitcomb, 
FSCA);  Pointe-a-Pitre,  Ilet  a  Boissard,  16°14'N, 
61°34'W,  26.vi.1960  (C.  &  R  Vaurie,  AMNH);  Terre- 
de-Haut,  Les  Saintes,  15°58'N,  61°35'W,  2.vii.l960 
(C.  &  P.  Vaurie,  AMNH).  Martinique:  Ansemitian  [?], 


Trois-Ilets,  14°33'N,  61°2'W,  10.vi.l960  (C.  &  P  Vau- 
rie, AMNH);  Diamant,  14°29'N,  61°2'W,  17.vi.l960 
(C.  &  P  Vaurie,  AMNH),  18.vi.l966  (C.  &  P  Vaurie, 
AMNH);  Fort  de  France,  14°36'N,  61°5'W, 
15.xii.l950  (N.  L.  H.  Krauss,  MCZ);  Pointe  Ferret, 
La  Caravelle,  14°45'N,  60°54'W,  19.vi.l960  (C.  &  R 
Vaurie,  AMNH);  Sainte-Anne,  14°26'N,  60°53'W, 
20.vi.l966  (C.  &  P  Vaurie,  AMNH);  Usine  de  Robert 
[?],  14°41'N,  60°57'W,  16.vi.l960  (C.  &  R  Vaurie, 
AMNH).  GRENADA  nr  Saint  Georges:  12°3'N, 
61°45'W,  3.vi.l950  (Leo  Isaacs,  AMNH).  NETH- 
ERLAND  ANTILLES  Bonaire:  Red  Pond  [?], 
12°12'N,  68°15'W,  3.i.l968  (B.  Malkin,  AMNH).  Cu- 
ragao:  12°11'N,  68°58'W,  13. i. 1968  (B.  Malkin, 
AMNH);  Fuik  (Oostpunt),  muddauber  nests,  12°4'N, 
68°49'W,  20.xii.l962  (H.  W.  Levi  &  B.  de  Jong, 
MCZ);  Groot  Sint  Joris,  plantation,  12°14'N,  69°3'W, 
22.xii.1962  (H.  W.  Levi  &  B.  de  Jong,  MCZ);  Groote 
Berg,  12°11'N,  69°0'W,  19.xii.l962  (H.  W.  &  L.  Levi, 
MCZ);  Hato,  12°11'N,  68°58'W,  28.xii.1967  (B.  Mal- 
kin, AMNH),  7.i.l968  (B.  Malkin,  AMNH);  Piscadera 
Baai,  12°8'N,  68°59'W,  18.xii.l962  (H.  W.  Levi, 
MCZ),  20.xii.l962  (H.  W.  Levi,  MCZ);  Savonet; 
shady  ravine,  12°20'N,  69°7'W,  28.xii.1962  (H.  W. 
Levi,  MCZ);  SE  of  airport,  12°10'N,  68°54'W, 
20.xii.l962  (H.  W.  Levi  &  B.  de  Jong,  MCZ);  Siberie, 
12°14'N,  69°3'W,  25.xii.1962  (H.  W.  Levi,  MCZ);  Sint 
Jan,  12°15'N,  69°6'W,  25.xii.1962  (H.  W.  Levi  &  B. 
de  Jong,  MCZ);  Willemstad,  Jewish  Cemetary, 
12°7'N,  68°57'W,  24.xii.I962  (H.  W.  Levi,  MCZ).  Sint 
Eustatius:  Oranjestad,  17°29'N,  62°59'W,  18.i.l968 
(B.  Malkin,  AMNH).  Sint  Maarten:  nr.  Juliana  Air- 
port [?],  18°4'N,  63°4'W,  24.ii.1965  (H.  Heatwole  & 
F.  MacKenzie,  AMNH).  PARAGUAY  Alto  Parana: 
Taguarazaya  [?],  25°30'S,  54°50'W  (AMNH).  PERU 
Ancash:  Huaraz,  9°32'S,  77°32'W,  6.xii.l980  (C.  Gold, 
CAS).  Apuriniac:  35  mi  E  Abancay,  13°38'S,  72°22'W, 
5.iii.l951  (E.  S.  Ross  &  Michelbacher,  CAS);  40  mi 
E  Abancay,  13°38'S,  72°20'W,  4.iii.l951  (E.  S.  Ross 
&  Michelbacher,  CAS);  Abancay,  13°38'2"S, 
72°52'53"W,  6.iii.l951  (E.  S.  Ross  &' Michelbacher, 
CAS).  Cajamarca:  Jaen,  5°42'34"S,  78°48'32"W, 
17.V.1967  (A.  F.  Archer,  AMNH).  Cusco:  40  mi  W 
Cusco,  13°32'S,  72°33'W,  5.iii.l951  (E.  S.  Ross  & 
Michelbacher,  CAS);  Hacienda  Urco,  near  Galea, 
13°22'S,  71°54'W,  19. ix.  1939  (Karl  P  Schmidt, 
AMNH);  Huacerpay  [?],  13°37'S,  72°13'W,  10.ix.l993 
(J.  Ochoa  Camara,  MCZ);  Ollantaitambo,  13°15'17"S, 
72°15'48"W,  15.vii.l911  (Yale  Pemvian  Expedition, 
AMNH),  15.xii.l980  (C.  Gold,  CAS);  Pisac, 
13°25'21"S,  71°50'48'W,  13.-xii.l980  (C.  Gold,  CAS); 
Urubamba,  13°18'28"S,  72°6'55"W,  15.i.l965  (Carras- 
co,  MCZ),  18.ii.l965  (H.  W.  Levi,  MCZ),  6.viii.l987 
(D.  Silva,  MUSM).  Huanuco:  Huanuco,  9°55'S, 
76°14'W,  6.X.1946  (J.  C.  Palhster,  AMNH).  lea:  km 
367  between  lea  and  Nazca,  sandy  semidesert, 
14°24'S,  75°23'W,  22.i.l952  (W.  Weyrauch,  CAS); 
Nazca,  14°50'S,  74°57'W,  15. iv.  1951  (P.  Aguilar, 
GAS).  Piura:  Gerro  Prieto,  La  Brea,  4°41'S,  81°6'W 
(CAS);  Mallares,  4°51'S,  80°46'W,  8.vi.l941  (H.  E.  F. 
&  D.  E.  F,  CAS),  13.vii.l941  (H.  E.  F.  &  D.  E.  F, 


52         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


CAS);  N  of  Negritos,  Farinas  Valley,  4°41'S,  81°18'W, 
9.X.1938  (D.  L.  &  H.  E.  Frizzell,  CAS);  nr.  Negritos, 
4°42'S,  81°18'W,  5.m.l939  (H.  E.  F,  CAS);  nr.  Se- 
chura,  5°33'S,  80°51'W,  4.xi.l941  (D.  L.  F,  CAS);  Ne- 
gritos, 4°42'S,  81°18'W,  15.xi.l934  (H.  E.  F,  CAS), 
15.iii.l941  (H.  E.  F,  CAS);  Farinas,  4°30'S,  81°8'W, 
7.V.1939  (D.  L.  &  H.  E.  Frizzell,  CAS),  31.V.1939  (D. 
L.  &  H.  E.  Frizzell,  CAS),  15.X.1939  (D.  L.  &  H.  E. 
Frizzell,  CAS);  Farinas  Valley,  4°30'S,  81°8'W, 
6.iii.l939  (D.  L.  &  H.  E.  Frizzell,  CAS),  8.iv.l939  (D. 
L.  &  H.  E.  Frizzell,  CAS);  Fortachuelo,  4°59'S, 
79°54'W,  29.iv.1939  (M.  H.  &  H.  E.  Frizzell,  CAS); 
Quebrada  Mogollon,  4°32'S,  81°4'W,  30.iv.l939  (D. 
L.  &  H.  E.  Frizzell,  CAS),  ll.vi.l939  (D.  L.  &  H.  E. 
Frizzell,  CAS);  Rio  Chira  Valley,  N  of  Amotape, 
4°53'S,  81°1'W,  13,xi.l938  (D.  L.  &  H.  E.  Frizzell, 
CAS);  S  of  Sechura,  5°33'S,  80°51'W,  25.X.1941  (D. 
L.  F,  CAS);  San  Lorenzo  [?],  Zona  Alta  Este  Herrera, 
5°4'S,  79°47'W,  12.iv.l969  (F  Aguilar,  MCZ);  Sullana, 
4°53'S,  80°41'W,  8.X.1939  (D.  L.  &  H.  E.  Frizzell, 
CAS),  5.X.1941  (D.  L.  &  H.  E.  Frizzell,  CAS).  Tum- 
bes:  34  km  E,  25  km  N  Funta  Farinas,  4°18'S, 
80°51'W,  l.i.l939  (D.  L.  &  H.  E.  Frizzell,  CAS). 
SAINT  LUCIA  Castries:  14°1'N,  61°0'W,  28.vii.1963 
(E.  N.  Kjellesvig-Waering,  AMNH).  TRINIDAD  & 
TORAGO  Tohago  Island:  Ruccoo  Ray,  11°10'N, 
60°48'W,  15.viii.l965  (E.  N.  Kjellesvig-Waering, 
AMNH);  Guayaguayare  Foint,  10°8'N,  61°2'W, 
14.ix.l963  (E.  N.  Kjellesvig-Waering,  AMNH);  Pi- 
geon Foint,  11°10'N,  60°50'W,  18.viii.l937  (E.  D., 
MCZ);  Salybia  Ray,  by  shore,  10°42'N,  61°2'W, 
15.ii.l972  (J.  A.  L.  Cooke,  MCZ);  Toco,  10°50'N, 
60°57'W,  19. iv.  1964  (Erik  N.  Kjellesvig-Waering, 
AMNH).  Trinidad:  Gasparee,  10°46'N,  61°19'W, 
3.xi.l944  (R.  Montgomery,  AMNH).  URUGUAY  Co- 
lonia:  Fmita  Gorda  [?],  34°28'S,  57°51'W,  25.ii.1968 
(R.  Capocasale  &  L.  Rruno,  CAS).  USA  Puerto  Rico: 
Algodones  Key,  18°12'N,  65°41'W,  15.x.  1964  (H. 
Heatwole,  R.  Levins  &  F  MacKenzie,  AMNH);  Ral- 
neario  [?]  Guajataca,  18°21'N,  66°55'W,  4.vii.l958  (A. 
F  Archer,  AMNH);  Railos  de  Coamo,  17°59'N, 
67°3'W,  2.iv.l990  (H.  W  &  L.  Levi,  MCZ),  3.iv.l990 
(H.  W  &  L.  Levi,  MCZ);  Rarranquitas,  18°11'N, 
66°18'W,  28.xii.1977  (J.  Coddington,  USNM);  below 
Quebradillas  along  old  RR  track,  18°28'N,  66°56'W, 
30.iii.l989  (H.  W  &  L.  Levi,  MCZ);  Cabeza  de  Ferro 
Island,  18°15'N,  65°35'W,  16.i.l965  (H.  Heatv^^ole  & 
F.  MacKenzie,  AMNH);  Cayo  Don  Luis,  17°57'N, 
66°58'W  12.i.l966  (Island  Froject  Staff,  Univ.  of 
Fuerto  Rico,  AMNH);  Cayo  Norte,  off  Culebra, 
18°20'N,  65°15'W,  14.ivl965  (H.  Heatwole  &  F. 
MacKenzie,  AMNH);  Cayo  Pinerito,  18°15'N, 
65°36'W,  24.ix.1964  (H.  Heatwole  &  F.  MacKenzie, 
AMNH);  Channel  at  Culebra,  Isla  del  Diablo, 
18°23'N,  65°40'W,  12.viii.l965  (Island  Froject  Staff, 
Univ.  of  Fuerto  Rico,  AMNH);  Cuevas  de  los  Alfaros, 
Rarrio  Mora,  18°29'N,  67°1'W,  20.vii.l958  (A.  F.  Ar- 
cher &  Rolle,  AMNH);  Culebra  Island,  18°19'N, 
65°17'W,  19.vii.l965  (F.  MacKenzie,  AMNH);  Cule- 
bra, near  Dewey,  18°18'N,  65°18'W,  10.viii.l965  (Is- 
land Froject  Staff,  Univ.  of  Fuerto  Rico,  AMNH);  Cu- 


lebrita  Island,  18°19'N,  65°14'W,  15.ivl965  (H.  Hea- 
twole &  F  MacKenzie,  AMNH),  ll.viii.l965  (Island 
Froject  Staff,  Univ.  of  Fuerto  Rico,  AMNH);  Dese- 
cheo  Is,  18°23'N,  67°29'W,  19.ii.l914  (AMNH);  De- 
secheo  Island,  18°23'N,  67°29'W,  28.V.1965  (H.  Hea- 
twole, R.  Levins  &  F  MacKenzie,  AMNH);  Faro  de 
Cabo  Rojo,  18°5'N,  67°9'W,  13.iii.l961  (F.  Rolle, 
AMNH);  Frank  Key,  Isl.  #13  nr.  La  Farguera  area, 
17°58'N,  67°3'W,  14.i.l966  (Island  Froject  Staff, 
Univ.  of  Fuerto  Rico,  MCZ);  Heatwole  Island,  off 
Culebrita,  18°19'N,  65°13'W,  14.iv.l965  (H.  Heatwo- 
le &  F.  MacKenzie,  AMNH);  Hormiqueros,  18°9'N, 
67°8'W,  16.ii.l962  (Aida  Velez,  AMNH),  ll.iii.l962 
(Aida  Velez,  AMNH),  19.iii.l962  (Aida  Velez, 
AMNH);  Isabela,  subexperiment  station,  18°30'N, 
67°1'W,  17.viii.l957  (A.  F.  Archer,  AMNH);  Jayuya, 
coffee  plantation,  18°13'N,  66°37'W,  23.iii.1986  (H. 
W.  &  L.  Levi,  MCZ);  Juana  Diaz,  18°3'N,  66°31'W, 
7.xi.l971  (J.  E.  Carico,  JEC);  Levins  Rock  [?], 
18°12'N,  65°41'W,  15.X.1964  (H.  Heatwole,  R.  Levins 
&  F.  MacKenzie,  AMNH);  Loma  Tinaja  [?],  S  of  La- 
guna  Cartagena,  17°59'N,  67°6'W  5.vii.l958  (A.  F. 
Archer  &  M.  Sanderson,  AMNH);  Luquillo,  near 
beach,  18°23'N,  65°44'W,  24.i.l932  (A.  S.  Mills, 
AMNH),  24.xii.1985  (V.  &  R.  Roth,  CAS),  27.iii.1988 
(H.  W  &  L.  Levi,  MCZ);  Mayagiiez,  18°12'N, 
67°9'W,  15.vii.l958  (A.  F.  Archer,  MCZ);  Mayagiiez, 
5  km  N  university  campus,  18°12'N,  67°9'W,  5.i.l964 
(MCZ);  Mayagiiez:  Las  Mesas,  18°11'N,  67°6'W, 
20.xi.l960  (F.  Rolle,  AMNH);  Mayagiiez,  university 
farm  N  university  campus,  18°12'N,  67°9'W,  7.ii.l964 
(MCZ);  McKenzie  Key,  Key  #3  [?],  18°12'N, 
65°41'W,  29.X.1964  (H.  Heatwole,  R.  Levins  &  F. 
MacKenzie,  AMNH);  Mona  Island,  Serralles,  18°5'N, 
67°54'W  [?],  7.ivl944  (Reatty,  MCZ);  Muertos  Is- 
land, beating  bushes,  17°54'N,  66°32'W  28.vl959 
(Jordan  &  Martorell,  AMNH);  Muertos  Island,  mud) 
nests  of  Sceliphron  caementarium,  17°54'N,  66°32'W, 
27.V.1959  (Medina  &  Martorell,  AMNH);  N  slope  Ti- 
naja, nr.  Cartagena  Lagoon,  beating  and  sweeping, 
18°23'N,  67°10'W,  5.vii.l958  (M.  W  Sanderson, 
AMNH);  Palominitos,  18°20'N,  65°34'W,  16.vi.l965 
(Island  Froject  Staff,  Univ.  of  Fuerto  Rico,  AMNH); 
Falomino  Island,  18°21'N,  65°34'W,  7.xi.l964  (H. 
Heatwole  &  F.  MacKenzie,  AMNH);  Farguera, 
17°59'N,  67°3'W,  25.iii.1990  (H.  W  &  L.  Levi,  MCZ); 
Fatillas,  18°0'N,  66°1'W,  3.iv.l931  (Mills  &  Leonard, 
AMNH);  Fico  Atalaya,  Rte.  2,  nr  Aiiasco,  18°18'N, 
67°11'W,  3.vii.l958  (M.  W.  Sanderson,  AMNH);  Fi- 
fieros  Island,  18°15'N,  65°35'W,  24.ix.1964  (H.  Hea- 
twole &  F.  MacKenzie,  AMNH);  Flaya  de  Humacao, 
18°10'N,  65°45'W,  23.V.1964  (MCZ);  Quebradillas, 
around  hotel,  18°29'N,  66°56'W,  30.iii.l989  (H.  W.  & 
L.  Levi,  MCZ);  Ratones  Island,  17°56'N,  66°17'W, 
19.xi.l964  (H.  Heatwole  &  R.  Levins,  AMNH);  Rom- 
ero III  [?],  17°57'N,  67°0'W,  13.i.l966  (Island  Froject 
Staff,  Univ.  of  Fuerto  Rico,  AMNH);  Rubianes, 
Caimito  Rajo  [?],  Cord.  Jaicoa,  18°26'N,  67°8'W, 
19.vii.l958  (A.  F  Archer  &  Rolle,  AMNH);  Santurce, 
18°27'N,  66°4'W,  28.iii.1931  (A.  S.  Mills,  AMNH); 
South  of  Corozo,  Cabo  Rojo  salt  flats,  edge  of  salt 


Metepeira  •  Piel 


53 


ponds,  17°56'N,  67°11'W,  23.iii.1990  (H.  W.  &  L. 
Levi,  MCZ);  Valle  de  Lajas,  18°1'N,  67°8'W,  3.vi.l958 
(A.  F.  Archer,  AMNH);  xerie  hills  N  of  Guanica, 
18°0'N,  66°55'W,  28.viii.1957  (A.  F.  Archer,  AMNH); 
Zancudo  Island  (Isleta  Marina),  18°20'N,  65°37'W, 
2..xi.l964  (H.  Heatwole  &  F.  MacKenzie,  AMNH). 
U.S.  VIRGIN  ISLANDS  Big  Cockroach:  18°24'N, 
65°4'W,  7.vi.l966  (Island  Project  Staff,  Univ.  of 
Puerto  Rico,  AMNH).  Congo  Cay:  18°22'N,  64°48'W, 
12.xi.l966  (Island  Project  Staff,  Univ.  of  Puerto  Rico, 
AMNH).  Grass  Cay:  18°22'N,  64°50'W,  12.xi.l966 
(Univ.  of  Puerto  Rico,  AMNH).  Great  Saint  James 
Island:  18°19'N,  64°50'W,  13.xi.l966  (Island  Project 
Staff,  Univ.  of  Puerto  Rico,  AMNH).  Hans  Lollik  Is- 
land: 18°24'N,  64°55'W,  6.iv.l966  (Island  Project 
Staff,  Univ.  of  Puerto  Rico,  AMNH).  Little  Cock- 
roaches: 18°25'N,  65°3'W,  7.vi.l966  (Island  Project 
Staff,  Univ.  of  Puerto  Rico,  AMNH).  Little  Hans  Lol- 
lick:  18°25'N,  64°54'W,  5.iv.l966  (Island  Project  Staff, 
Univ.  of  Puerto  Rico,  AMNH).  Little  Saint  James  Is- 
land: 18°18'N,  64°50'W,  13.xi.l966  (Island  Project 
Staff,  Univ.  of  Puerto  Rico,  AMNH).  Mingo  Cay: 
18°22'N,  64°49'W,  12.xi.l966  (Island  Project  Staff, 
Univ.  of  Puerto  Rico,  AMNH).  Saint  John:  18°20'N, 
64°45'W,  9.iii.l925  (F.  E.  Lutz,  AMNH),  ll.vii.l958 
(A.  F  Archer  &  M.  Sanderson,  AMNH),  15.xii.l967 
(N.  L.  H.  Krauss,  AMNH);  Cruz  Ray,  18°20'N, 
64°48'W,  27.ii.1964  (A.  M.  Chickering,  MCZ);  nr. 
Cinnamon  &  Hart  Bays  on  W  half  of  island,  18°20'N, 
64°46'W,  2.vlii.l976  (D.  E.  &  D.  N,  Rosen,  AMNH). 
Saint  Thomas:  18°24'N,  64°55'W,  24. ii. 1925 
(AMNH),  24.vii.1925  (A.  Petrunkevitch,  CAS), 
24.xi.1925  (AMNH);  Crov^^n  Mountain,  18°21'N, 
64°58'W,  30.viii.l957  (A.  F.  Archer,  AMNH), 
7.vii.l958  (A.  F.  Archer,  AMNH);  Denmark  Hill  [?], 
18°24'N,  66°55'W,  l.ix.l957  (A.  F  Archer,  AMNH); 
Flagstok  Hill,  Stumpy  Bay,  18°22'N,  65°0'W, 
9.vii.l958  (A.  F.  Archer,  AMNH);  Harmans,  Char- 
lotte Amalia,  18°21'N,  64°56'W,  2.ix.l957  (A.  F.  Ar- 
cher &  family,  AMNH);  Hassell  Island,  18°20'N, 
64°56'W,  l.ix.l957  (A.  F  Archer,  AMNH),  15.ii.l964 
(A.  M.  Chickering,  MCZ),  10.viii.l966  (M.  L.  Pres- 
sick,  AMNH).  St.  Croix:  Christiansted,  17°45'N, 
64°42'W,  3.vi.l911  (AMNH),  15.i.l955  (A.  M.  Nad- 
ler,  AMNH);  East  End,  17°45'N,  64°40'W,  15.xii.l965 
(Island  Project  Staff,  Univ.  of  Puerto  Rico,  AMNH); 
Green  Key,  17°46'N,  64°40'W,  17.iv.l964  (H.  Hea- 
twole, MCZ);  Mount  Eagle,  dryish  forest,  17°46'N, 
64°49'W,  15.xii.l965  (Island  Project  Staff,  Univ.  of 
Puerto  Rico,  AMNH);  Protestant  Cay,  17°45'N, 
64°42'W,  18.iv.l964  (H.  Heatwole,  MCZ);  St.  Croix, 
17°45'N,  64°54'W,  l.ix.l966  (Chickering,  MCZ), 
6.ix.l966  (Chickering,  MCZ).  VENEZUELA  Depen- 
dencias  Federales:  Patos,  10°38'N,  61°52'W, 
23.ix.1944  (R.  H.  Montgomery,  AMNH).  Districto 
Federal:  Punta  Tanaguarena,  in  coastal  bldg.  &  gar- 
den, 10°37'N,  66°48'W,  26.xii.1970  (W.  B.  Peck, 
CAS). 


1 7.  Metepeira  roraima  new  species 
Figures  141-148,  316;  Map  8 

Holotype.  Female  from  Rio  Suioimu,  Roraima,  Brazil, 
X.1966,  M.  Abrorenga,  in  MZSP  The  specific  name 
is  a  noun  in  apposition  after  the  locality. 

Description.  Female  holotype.  Brown 
carapace;  lighter  around  eyes  (Fig.  146). 
Coxae,  femora,  tibiae,  and  patellae  tan, 
lighter  ventrally  Metatarsi,  tarsi  white.  Fe- 
mur I  with  row  of  three  or  four  macrose- 
tae  on  anterior  side;  none  on  anteroventral 
side.  Dorsal  folium  with  typical  Metepeira 
fleur-de-lis  pattern  (Fig.  146).  Venter  of 
abdomen  brown  with  wide  white  median 
line,  flanked  by  thinner  white  lines  that  to- 
gether form  a  T-shape  posteriorly  (Fig. 
147).  Pair  of  small  white  spots  on  either 
side  of  T-shape  mark.  Sternum  brown  with 
median  white  line  (Fig.  147).  Ratio  of  eye 
diameters:  posterior  medians  and  anterior 
medians  1.1,  anterior  laterals  1.5,  posterior 
laterals  1.3.  Anterior  median  eyes  separat- 
ed by  1.2  diameters,  posterior  median  eyes 
by  0.6,  anterior  median  eyes  separated 
from  anterior  laterals  by  1.9  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.1  their  diameters.  Total  length  4  mm. 
Carapace  1.8  mm  long,  1.4  wide.  First  fe- 
mur 1.9  mm,  patella  and  tibia  2.1,  meta- 
tarsus 1.8,  tarsus  0.7.  Second  patella  and 
tibia  1.7  mm,  third  1.0,  fourth  1.6. 

Male  paratype  from  Rio  Surumu,  Ro- 
raima, Brazil.  Carapace  brown;  lighter 
around  eyes  with  median  white  mark  (Fig. 
144).  Legs  brown,  white  at  base  of  femora. 
Femur  I  with  row  of  four  to  five  macro- 
setae  on  anterior  side;  four  to  five  on  an- 
teroventral side.  Dorsal  folium,  venter,  and 
sternum  as  in  female  (Figs.  144,  145).  Ra- 
tio of  eye  diameters:  posterior  medians 
and  anterior  medians  1.1,  anterior  laterals 
1.6,  posterior  laterals  1.4.  Anterior  median 
eyes  separated  by  1.4  diameters,  posterior 
median  eyes  by  0.6,  anterior  median  eyes 
separated  from  anterior  laterals  by  1.6  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.2  their  diameters.  To- 
tal length  2.9  mm.  Carapace  1.3  mm  long, 
1.1  wide.  First  femur  1.9  mm,  patella  and 


54         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


tibia  2,  metatarsus  1.6,  tarsus  0.6.  Second 
patella  and  tibia  1.5  mm,  third  0.8,  fourth 
1.3. 

Diagnosis.  The  epigynum  of  M.  roraima 
differs  from  that  of  M.  compsa  by  having 
oval  openings  (Fig.  143)  instead  of  round 
openings  (Figs.  131,  134);  it  differs  from 
M.  gressa  by  being  more  translucent  and 
by  having  a  narrower  scape  (compare  Fig. 
143  with  Fig.  151).  Unlike  M.  gres.sa,  the 
embolus  tip  of  M.  roraima  is  more  bent 
and  the  distal  apophysis  is  as  wide  as  the 
embolus  tip  (compare  Fig.  141  with  Fig. 
149).  Unlike  M.  compsa,  the  distal  embolic 
apophysis  projects  away  from  the  embolus 
tip  (Fig.  141)  instead  of  extending  straight 
from  the  base  and  parallel  to  the  embolus 
tip  (Figs.  129,  132). 

Variation.  Average  body  length  of  eight 
females  examined  5  mm,  range  4  to  7  mm. 
Average  body  length  of  five  males  exam- 
ined 3.1  mm,  range  2.8  to  4  mm.  Colom- 
bian specimens  are  much  darker  than 
those  from  northern  Brazil  and  southern 
Guyana,  suggesting  the  possibility  that 
these  populations  represent  separate  cryp- 
tic species.  In  fact,  the  carapace  on  Col- 
ombian specimens  is  often  jet  black  with 
the  white  marks  on  the  dorsal  folium 
much  reduced  (Fig.  148).  However,  there 
is  little  corresponding  genitalic  difference, 
especially  among  males;  consequently, 
these  populations  are  deemed  to  be  con- 
specific. 

Natural  History.  Mature  M.  roraima 
specimens  from  eastern  South  America 
have  been  collected  in  November  and  De- 
cember; specimens  from  western  South 
America  have  been  collected  between 
March  and  July  (Fig.  316).  Median  eleva- 
tion, 1,000  m. 

Distribution.  Western  Colombia,  north- 
ern Brazil,  and  southern  Guyana  (Map  8). 

Records  Examined.  BRAZIL  Roraima:  Rio  Suru- 
mu,  Jerrit  Rio  Branco  [?],  3°22'N,  60°19'W,  x.1966 
(M.  Abrorenga,  MZSP).  COLOMBIA.  Valle  del  Cau- 
ca:  Atuncela,  3°46'N,  76°42'W,  19.vii.l970  (W.  Eber- 
hard,  MCZ);  Cali,  around  house,  3°27'N,  76°31'W 
(W.  Eberhard,  MCZ);  Palmira,  3°32'N,  76°17'W, 
l.iii.l964  (Ballo,  CAS);  Rio  Panee,  below  Buenos  Ai- 


res, 3°55'N,  76°8'W,  5.iv.l970  (W.  Eberhard,  MCZ); 
Rio  Pance,  near  Cali.,  3°21'N,  76°33'W,  8.V.1970  (W. 
Eberhard,  MCZ),  8.vi.l970  (W.  Eberhard,  MCZ), 
15.vi.l970  (W.  Eberhard,  MCZ),  23.vi.1970  (W. 
Eberhard,  MCZ),  25.vi,1973  (W.  Eberhard,  MCZ). 
GUYANA  Upper  Takiitu:  Isherton,  on  lat.  2,  10  mi  E 
Rupununi  River,  2°19'N,  59°22'W,  15.xi.l937  (W.  G. 
Hassler,  AMNH). 

18.  Metepeira  gressa  (Keyserling) 
Figures  149-156,  300;  Map  8 

Epeira  gressa  Keyserling,  1892:  166,  fig.  123,  9 .  Five 
female  syntypes  from  Taquara,  Rio  Grande  do  Sul, 
Brazil,  in  BMNH,  examined.  One  specimen  des- 
ignated lectotype. 

Epeira  seditiosa  Keyserling,  1893:  212,  fig.  157,  S. 
Male  holotype  from  Rio  Grande  do  Sul,  Brazil,  in 
BMNH,  examined.  NEW  SYNONYMY. 

Araneus  gressus: — Petrunkevitch,  1911:  314.  Roewer, 
1942:  844.  Bonnet,  1955:  511. 

Araneus  seditiosus: — Petrunkevitch,  1911:  314. 
Roewer,  1942:  852.  Bonnet  (1955:  592)  erroneously 
suggests  that  Petrunkevitch  (1911:  314)  synony- 
mized  Araneus  scitulus  (Blackwall,  1863)  with  Ar- 
aneus seditiosus. 

Eustala  seditiosa: — Mello-Leitao,  1943:  179.  Erro- 
neous transfer. 

Metazygia  gressa: — Mello-Leitao,  1943:  187.  Erro- 
neous transfer. 

Metepeira  gressa: — Levi,  1991:  179.  Platnick,  1993: 
449. 

Metepeira  seditiosa: — Levi,  1991:  180.  Platnick,  1993: 
449. 

Note.  Examination  of  voucher  specimens  of  Vi- 
era  suggest  that  in  the  behavioral  studies  of  Viera, 
(1986,  1989)  and  Viera  and  Costa  (1988),  tlie  name! 
"Metepeira  sp.  A"  is,  in  fact,  M.  gressa. 

Description.  Female  from  Punta  del  Es- 
pinillo,  Montevideo,  Uruguay.  Chelicerae 
brown,  lighter  on  distal  inside  margins. 
Carapace  brown;  yellowish  white  across 
and  just  behind  posterior  eye  row;  median 
white  line  reaching  thoracic  furrow,  some- 
times thickened  into  arrow  shape  (Fig. 
155).  Proximal  halves  of  femora,  white;  re- 
mainder black.  Patellae  black  ventrally, 
white  dorsally.  Distal  halves  of  tibiae, 
black;  remainder  white  with  black  annu- 
lation.  Femur  I  with  row  of  four  macro- 
setae  on  anterior  side;  none  on  antero- 
ventral  side.  Dorsal  folium  white  fleur-de- 
lis  pattern  on  black,  margined  with  wavy 
white  lines  (Fig.  155).  Venter  black  with 
short,  wide,  white  median  line;  pair  of 
small  white  spots  on  either  side  of  spiracle 


Metepeira  •  Piel        55 


(Fig.  156).  Sternum  black  with  median 
white  hne,  often  broken  (Fig.  156).  Ratio 
of  eye  diameters:  posterior  medians  and 
anterior  medians  1.2,  anterior  laterals  1.3, 
posterior  laterals  1.3.  Anterior  median 
eyes  separated  by  1.8  diameters,  posterior 
median  eyes  by  1.0,  anterior  median  eyes 
separated  from  anterior  laterals  by  2  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.2  their  diameters.  To- 
tal length  3.3  mm.  Carapace  1.8  mm  long, 
1.4  wide.  First  femur  1.9  mm,  patella  and 
tibia  2,  metatarsus  1.6,  tarsus  0.7.  Second 
patella  and  tibia  1.8  mm,  third  1.1,  fourth 
1.6. 

Male  from  Punta  del  Espinillo,  Monte- 
video, Uruguay.  Coloration  of  chelicerae 
and  carapace  as  in  female  except  median 
line  brighter,  thickened  into  arrow  shape 
(Fig.  153).  Leg  coloration  as  in  female.  Fe- 
mur I  with  row  of  four  macrosetae  on  an- 
terior side;  five  to  six  on  anteroventral  side. 
Dorsal  folium,  venter,  and  sternum  as  in 
female  (Figs.  153,  154).  Ratio  of  eye  di- 
ameters: posterior  medians  and  anterior 
medians  1.0,  anterior  laterals  1.5,  posterior 
laterals  1.4.  Anterior  median  eyes  separat- 
ed by  1.3  diameters,  posterior  median  eyes 
by  0.8,  anterior  median  eyes  separated 
from  anterior  laterals  by  1.9  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.3  their  diameters.  Total  length  3  mm. 
Carapace  1.8  mm  long,  1.3  wide.  First  fe- 
mur 2.5  mm,  patella  and  tibia  2.3,  meta- 
tarsus 1.9,  tarsus  0.9.  Second  patella  and 
tibia  1.8  mm,  third  1.1,  fourth  1.6. 

Diagnosis.  The  epigynum  of  M.  gressa 
differs  from  that  of  M.  compso  by  having 
oval  openings  (Fig.  151)  instead  of  round 
openings  (Figs.  131,  134);  it  differs  from 
M.  roraima  by  being  less  translucent  and 
by  having  a  wider  scape  (compare  Fig.  151 
with  Fig.  143).  Unlike  M.  roraima,  the  em- 
bolus tip  in  M.  gressa  is  not  as  bent,  and 
the  distal  apophysis  is  thinner  than  the 
point  on  the  embolus  where  it  is  attached 
(compare  Fig.  149  with  Fig.  141).  Unlike 
M.  compsa,  the  distal  embolic  apophysis 
projects  away  from  the  embolus  tip  (Fig. 
149)  instead  of  extending  straight  from  the 


base  and  parallel  to  the  embolus  tip  (Figs. 
129,  132). 

Variation.  Average  body  length  of  14  fe- 
males examined  4.8  mm,  range  3.3  to  6.8 
mm.  Average  body  length  of  five  males  ex- 
amined 3.4  mm,  range  2.8  to  4.3  mm. 

Natural  History.  In  Uruguay,  M.  gressa 
live  in  small  colonies  of  up  to  five  individ- 
uals surrounding  the  inflorescences  of 
Eryngium  sp.  (Viera  and  Costa,  1988).  The 
number  of  sticky  spirals  (14  below  the 
hub,  22  above  the  hub)  and  radii  (c.  40) 
are  the  same  irrespective  of  age;  however 
the  length  and  width  of  the  web  differ  be- 
tween juveniles  (c.  6  and  5  cm,  respective- 
ly) and  adults  (c.  9  and  7  cm,  respectively) 
(Viera,  1992).  Mature  specimens  are  most 
often  collected  between  September  and 
March  at  low  elevation  (Fig.  300). 

Distribution.  Northern  Argentina,  Par- 
aguay, and  Uruguay  (Map  8). 

Records  Examined.  ARGENTINA  Buenos  Aires: 
Arrecifes,  34°3'S,  60°7'W,  17.1.1939  (Biraben,  MLP); 
Boulogne,  34°30'S,  58°34'W,  15.x. 1938  (Prosen, 
MLP);  Colon  [?],  33°53'29"S,  61°6'35"W,  20.Lx.l944 
(Torres,  MLP);  Moreno,  34°39'S,  58°48'W,  15.ii.l966 
(Rossi  and  Maury,  MACN);  San  Isidro,  34°27'S, 
58°30'W,  15.xii.l937  (Peregra,  MACN);  San  Mi- 
guel—San Fernando,  34°29'S,  58°39'W,  15.vii.l940 
(F.  Morrios,  MACN);  Sierra  de  la  Ventana,  38°9'S, 
61°48'W,  15.iii.l939  (S.  H.  Bavio,  MACN),  15.xi.l954 
(Fritz,  MACN),  31.X.1969  (Carlos  Grisolia,  MCZ), 
15.vii.l972  (Amarrilla,  MACN);  Tigre,  34°25'S, 
58°34'W  (J.  M.  Viana,  MACN).  Entre  Rios:  Concep- 
cion  del  Uruguay,  32°29'0"S,  58°13'42"W,  4.i.l941 
(Prosen,  MLP);  Salto  Grande,  31°13'S,  57°56'W, 
15.iii.l964  (M.  E.  Galiano,  MEG).  Santa  Fe:  Floren- 
cia  Varelo  [?],  28°2'S,  59°15'W,  15.xii.l939  (F.  Morris, 
MACN).  Tucumdn:  30  km  S  Concepcion,  27°36'S, 
65°35'W,  16.i.l983  (L.  E.  Pena,  AMNH).  PARA- 
GUAY Itapiia:  San  Luis,  27°6'S,  56°36'W,  15.X.1908 
(AMNH).  URUGUAY  Canelones:  Montevideo:  De- 
tras  del  Cerro,  Camino  de  las  tropas  [?],  34°45'S, 
56°10'W,  6.ii.l963  (R.  Capocasale  and  L.  Bmno, 
CAS).  Colonia:  Punta  Gorda^?],  34°28'S,  57°51'W, 
25. ii.  1968  (R.  Capocasale  and  L.  Bruno,  CAS), 
26.ii.1968  (R.  Capocasale  and  L.  Bmno,  CAS).  Mal- 
donado:  Colonia,  Cerro  de  las  Animas.  Small  stones 
in  native  forest,  34°46'S,  55°19'W,  30.x.  1967  (P  San 
Martin,  MCZ).  Montevideo:  Punta  del  Espinillo, 
34°50'S,  56°26'W,  12.X.1983  (Carmen  Viera,  MCZ), 
15.xii.l983  (Carmen  Viera,  CV).  Treinta  y  Tres:  Que- 
brada  de  los  Cuervos,  33°10'S,  54°27'W,  27.X.1990 
(Lopez,  Perez,  Viera,  MCZ). 


56         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


Metepeira  incrassata  Group 

Species  in  the  M.  incrassata  group  all 
lack  a  keel  on  the  median  apophysis.  Males 
have  a  distal  embolic  apophysis  that  either 
projects  toward  the  embolus  tip  (e.g..  Fig. 
157),  forms  a  shai-p  comer  but  does  not 
actually  project  forward  (e.g..  Fig.  185),  or 
forms  a  smoother,  rounded,  and  gradual 
comer  (e.g..  Figs.  192,  206).  The  M.  in- 
crassata species  group  includes  Metepeira 
maya,  Metepeira  inca,  Metepeira  gosoga, 
Metepeira  ohnec,  Metepeira  comanche, 
Metepeira  pimungan,  Metepeira  triangu- 
laris, Metepeira  arizonica,  Metepeira  atas- 
cadero,  and  Metepeira  incrassata. 

19.  Metepeira  maya  new  species 
Figures  157-163,  317;  IViap  14 

Holotype.  Male  from  North  Bay,  Twin  Cays,  Stann 
Creek  District,  Belize,  14.iii.l986,  P.  Sierwald,  in 
USNM.  The  specific  name  is  a  noun  in  apposition 
after  the  Indian  people  of  southern  Mexico  and 
Central  America. 

Description.  Female  paratype  from 
Twin  Cays,  Stann  Creek  District,  Belize. 
Carapace  brown  with  large  white  eye  re- 
gion, white  lateral  posterior  extensions, 
and  short  median  posterior  extension  (Fig. 
162).  Legs  white,  ringed  black  on  distal 
ends  of  articles.  Femur  I  with  row  of  four 
macrosetae  on  anterior  side;  none  on  an- 
te roventral  side.  Dorsum  of  abdomen  with 
usual  Metepeira  folium,  though  frequently 
with  deep  red  pigmentation  on  anterior 
half  (Fig.  162).  Venter  wide,  white  median 
line  with  pair  of  large  white  spots  on  either 
side  of  spiracle  (Fig.  163).  Sternum  with 
wide  median  white  line  widening  anteri- 
orly with  constriction  in  center  (Fig.  163). 
Ratio  of  eye  diameters:  posterior  medians 
and  anterior  medians  1.0,  anterior  laterals 
1.3,  posterior  laterals  1.3.  Anterior  median 
eyes  separated  by  1.2  diameters,  posterior 
median  eyes  by  0.8,  anterior  median  eyes 
separated  from  anterior  laterals  by  2  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.2  their  diameters.  To- 
tal length  3.9  mm.  Carapace  1.8  mm  long, 
1.4  wide.  First  femur  2.2  mm,  patella  and 
tibia  2.1,  metatarsus  1.6,  tarsus  0.8.  Sec- 


ond patella  and  tibia  1.8  mm,  third  1.0, 
fourth  1.6. 

Male  holotype.  Carapace  dark  with  light 
region  around  eyes  and  light  triangular 
mark  anterior  to  thoracic  furrow  (Fig. 
160).  Legs  darker  than  in  female.  Femur 
I  with  row  of  four  macrosetae  on  anterior 
side;  four  on  anteroventral  side.  Dorsum, 
venter,  sternum  as  in  female  except  me- 
dian white  line  on  sternum  usually  broken 
(Figs.  160,  161).  Ratio  of  eye  diameters: 
posterior  medians  and  anterior  medians 
1.0,  anterior  laterals  1.4,  posterior  laterals 
1.2.  Anterior  median  eyes  separated  by  1.2 
diameters,  posterior  median  eyes  by  0.9, 
anterior  median  eyes  separated  from  an- 
terior laterals  by  1.7  diameters  of  anterior 
lateral  eyes,  lateral  eyes  separated  by  0.3 
their  diameters.  Total  length  3  mm.  Car- 
apace 1.5  mm  long,  1.2  wide.  First  feiTiur 
2.2  mm,  patella  and  tibia  2.3,  metatarsus 
1.8,  tarsus  0.9.  Second  patella  and  tibia  1.8 
mm,  third  1.0,  fourth  1.4. 

Diagnosis.  Unlike  others  in  the  M.  in- 
crassata species  group,  M.  maya  and  M. 
inca  both  have  distal  embolic  apophyses 
that  project  foiAvard,  forming  rounded 
bumps  (Figs.  157,  164).  The  male  palps  of 
these  two  species  differ  by  the  thickness 
of  the  flagellae  and  base  on  the  median) 
apophysis:  thicker  in  M.  maya  than  in  M. 
inca  (compare  Fig.  157  with  Fig.  164).  The 
epigynum  of  M.  maya  resembles  those  of 
M.  inca  and  M.  comanche  because  the  oval 
epigynal  openings  are  formed  out  of  mem- 
branous surfaces  that  are  distinctly  sepa- 
rate from  the  base  of  the  scape  (Figs.  159, 
166,  187).  The  epigynal  openings  of  M. 
maya  (Fig.  159)  are  much  wider  than 
those  of  M.  comanche  (Fig.  187);  and  the 
posterior  lobes  on  M.  maya  are  thickened 
(Fig.  159)  but  not  pointed  as  in  M.  inca 
(Fig.  166). 

Variation.  Average  body  length  of  five 
females  examined  5.2  mm,  range  3.9  to  6.3 
mm.  Average  body  length  of  four  males 
examined  2.8  mm,  range  2.3  to  3.1  mm. 

Natural  History.  Mature  specimens  have 
been  collected  in   March  through  August 


Metepeira  •  Piel        57 


gressa 
(18) 


It 


maya 
(19) 


A 


r^;''i 


164 


169^^       170 

inca 
(20) 


Figures  149-156.  Metepeira  gressa  (Keyserling)  (sp.  18;  34°50'S,  56°26'W).  149,  male  palpus,  mesal.  150,  epigynum,  posterior. 

151,  epigynum,  ventral.  152,  male  embolic  division,  ventral.  153,  male,  dorsal.  154,  male,  ventral.  155,  female,  dorsal.  156, 

female,  ventral. 

Figures  157-163.  Metepeira  maya  new  species  (sp.  19;  16°50'N,  88°5'W).  157,  male  palpus,  mesak  158,  epigynum,  posterior. 

159,  epigynum,  ventral.  160,  male,  dorsal.  161,  male,  ventral.  162,  female,  dorsal.  163,  female,  ventral. 

Figures  164-170.  Metepeira  inca  new  species  (sp.  20;  4°30'S,  81°8'W).  164,  male  palpus,  mesal.  165,  epigynum,  posterior. 

166,  epigynum,  ventral.  167,  male,  dorsal.  168,  male,  ventral.  169,  female,  dorsal.  170,  female,  ventral. 

Scale  bars:  dorsum  and  venter  figures  1.0  mm. 


58         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


(Fig.  317).  Habitats  range  from  mangroves 
at  sea  level  to  pine  forests  at  1,600  m. 

Distribution.  Southern  Mexico  to  Costa 
Rica  (Map  14). 

Records  Examined.  BELIZE  Stann  Creek:  Twin 
Cays,  Andera  Flats,  16°50'N,  88°5'W,  20.iii.l986  (P. 
Sierwald,  USNM);  Twin  Cays,  North  part  of  West 
Pond,  West  of  Swamp  Doc,  16°50'N,  88°5'W 
28.iii.1986  (P  Siei-wald,  USNM);  Twin  Cays,  North- 
west Point,  North  Bay,  16°50'N,  88°5'W,  14.iii.l986 
(P.  Sieiw^ald,  USNM);  Twin  Cays,  Northwest  Point, 
North  Bay,  ground  not  flooded,  16°50'N,  88°5'W, 
14.iii.l986  (P  Sierwald,  USNM);  Twin  Cays,  red 
mangrove,  16°50'N,  88°5'W,  17.V.1985  (Feller, 
USNM);  Twin  Cays,  white  mangrove,  16°50'N, 
88°5'W,  15.vi.l984  (Feller,  USNM),  5.vi.l985  (Erwin, 
Mathis,  Sims,  USNM).  COSTA  RICA  San  Jose:  San 
Jose,  9°56'N,  84°5'W  (Tristan  &  Banks,  MCZ).  GUA- 
TEMALA Chiquimula:  Chiquimula,  14°48'N, 
89°33'W,  22.vii.1947  (C.  &  P  Vaurie,  AMNH).  Sa- 
catepequez:  Antigua,  14°34'N,  90°44'W,  16.viii.l947 
(C.  &  P  Vaurie,  AMNH).  MEXICO  Chiapas:  Com- 
itan  de  Dominguez,  16°1.5'N,  92°8'W,  19.vii.l950  (C. 
J.  &  M.  Goodnight,  AMNH);  near  Rio  San  Gregorio, 
between  Comitan  and  Ocotal,  15°45'N,  92°0'W, 
18.vii.l950  (C.  J.  &  M.  Goodnight,  AMNH);  pine  for- 
est, 15  mi  NW  Arriaga,  16°25'N,  94°1'W,  27.viii.1966 
(Jean  &  Wilton  Ivie,  AMNH);  Tuxda  Gutierrez, 
16°45'N,  93°7'W,  10.vi.l964  (Palhster,  AMNH).  NIC- 
ARAGUA Mfltogoipa;  Matagaslpa,  12°53'N,  85°57'W 
15.vii.l989  (R.  Reinbold,  JMM). 

20.  Metepeira  inca  new  species 
Figures  164-170,  318;  Map  11 

Holotype.  Male  from  Pariiias  Valley,  Piura,  Peru, 
21.V.1939,  D.  L.  &  H.  E.  Frizzell  in  CAS.  The 
specific  name  is  a  noun  in  apposition  after  the  Que- 
chuan  people  who  once  luled  Peru. 

Description.  Female  paratype  from  Par- 
iiias Valley,  Piura,  Peru.  Light  reddish 
brown  carapace,  white  around  eyes,  very 
faint  light  mark  in  center  (Fig.  169).  Legs 
white;  slightly  darker  on  dorsal  surfaces. 
Femur  I  with  row  of  four  to  five  macro- 
setae  on  anterior  side;  none  on  anterov- 
entral  side.  Dorsal  folium  white;  darker  on 
shoulders  and  posteriorly  (Fig.  169).  Ven- 
ter grayish  browii  with  wide  white  median 
mark  inside  U-shaped  pattern;  pair  of 
white  spots  on  either  side  of  spiracle  (Fig. 
170).  Sternum  white  (Fig.  170).  Ratio  of 
eye  diameters:  posterior  medians  and  an- 
terior medians  0.9,  anterior  laterals  1.0, 
posterior    laterals    1.1.    Anterior    median 


eyes  separated  by  1.6  diameters,  posterior 
median  eyes  by  1.3,  anterior  median  eyes 
separated  from  anterior  laterals  by  2.5  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.3  their  diameters.  To- 
tal length  6.8  mm.  Carapace  2.9  mm  long, 
2.3  wide.  First  femur  3  mm,  patella  and 
tibia  3.3,  metatarsus  2.6,  tarsus  1.  Second 
patella  and  tibia  2.8  mm,  third  1.7,  fourth 
2.5. 

Male  holotype.  Light  reddish  brown 
carapace,  white  around  eyes,  faint  light  tri- 
angular mark  in  center  (Fig.  167).  Legs 
white,  save  for  light  reddish  brown  on  dor- 
sal surfaces  of  femora  and  patellae.  Femur 
I  with  row  of  three  macrosetae  on  anterior 
side;  two  to  three  on  anteroventral  side. 
Dorsal  folium  white  with  wavy  black  lines 
thickening  posteriorly  (Fig.  167).  Venter 
and  sternum  as  in  female  (Fig.  168).  Ratio 
of  eye  diameters:  posterior  inedians  and 
anterior  medians  1.0,  anterior  laterals  1.3, 
posterior  laterals  1.1.  Anterior  median 
eyes  separated  by  1.3  diameters,  posterior 
median  eyes  by  0.8,  anterior  median  eyes 
separated  from  anterior  laterals  by  1.9  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  almost  touching.  Total  length  3.7  mm. 
Carapace  1.9  mm  long,  1.4  wide.  First  fe- 
mur 2.9  mm,  patella  and  tibia  2.9,  meta-) 
tarsus  2.7,  tarsus  1.  Second  patella  and  tib- 
ia 2.4  mm,  third  1.3,  fourth  1.9. 

Diagnosis.  Unlike  others  in  the  M.  in- 
crassata  species  group,  M.  inca  and  M. 
niaija  both  have  distal  embolic  apophyses 
that  project  forward,  forming  rounded 
bumps  (Figs.  157,  164).  The  male  palps  of 
these  two  species  are  separated  by  the 
thinner  flagellae  and  base  on  the  median 
apophysis  of  M.  inca  (compare  Fig.  164 
with  Fig.  157).  The  epigynum  of  M.  inca 
resembles  those  of  M.  inaija  and  M.  coni- 
anche  because  the  oval  epigynal  openings 
are  formed  out  of  membranous  surfaces 
that  are  distinctly  separate  from  the  base 
of  the  scape  (Figs.  159,  166,  187).  The  epi- 
gynal openings  of  M.  inca  (Fig.  166)  are 
much  wider  than  those  of  M.  conianche 
(Fig.  187);  the  posterior  lobes  on  M.  inca 


Metepeira  •  Piel 


59 


are  pointed  (Fig.  166),  not  just  thickened, 
as  in  M.  may  a  (Fig.  159). 

Variation.  Average  body  length  of  six  fe- 
males examined  5.7  mm,  range  5.2  to  6.8 
mm.  Average  body  length  of  four  males 
examined  3.7  mm,  range  2.8  to  4.4  mm. 
Coloration  varies  significantly  among  lo- 
calities, especially  in  the  males,  where 
some  are  frequently  much  darker  than 
others. 

Natural  History.  Mature  specimens 
have  been  collected  in  April  through  Oc- 
tober (Fig.  318)  at  elevations  between  300 
and  600  m. 

Distribution.  Most  northern  tip  of  Peru 
(Map  11). 

Records  Examined.  PERU  Piiira:  12  mi  N  Man- 
cora,  4°0'S,  80°54'W,  ll.xii.l93S;  Parinas  Valley, 
4°30'S,  81°8'W,  3.iv.l939,  8.iv.l939,  16.iv.l939, 
7.V.1939,  21. V.  1939,  25.vi.1939,  3.vii.l939,  6.viii.l939, 
15.viii.l939,  15.X.1939;  Quebrada  de  Pariiias,  4°32'S, 
81°17'W,  14.iv.l939,  7.V.1939,  21.V.1939;  Quebrada 
MogoUon,  4°32'S,  81°4'W,  30.iv.l939,  ll.vi.l939, 
18.vi.l939,  21.vi.l939,  ll.vii.l939,  16.vii.l939, 
24.ix.1939  (all  records:  D.  L.  &  H.  E.  Frizzell,  CAS). 

21 .  Metepeira  gosoga 
Chamberlin  and  Ivie 

Figures  171-177,  322;  Map  9 

Metepeira  gosoga  Chamberlin  and  Ivie,  1935:  21  figs 
82-83,  9  .  Female  holotype  from  Pilot  Knob  Valley, 
Mohave  Desert,  California,  in  the  AMNH,  exam- 
ined. Roewer,  1942:  868.  Ronnet,  1957:  2820.  Levi, 
1977:  200,  figs.  28-36. 

Description.  Female  from  Baja  Califor- 
nia Norte,  Mexico.  Carapace  diiiy  yellow- 
ish brown,  lighter  anterior  half,  darker  di- 
amond-shaped mark  behind  eyes  (Fig. 
176).  Legs  same  color  as  carapace;  dark 
rings  on  distal  ends  of  articles.  Femur  I 
with  row  of  three  to  five  inacrosetae  on 
anterior  side;  two  or  three  macrosetae  on 
anteroventral  side.  Folium  lighter  than  in 
other  species,  darkening  posteriorly  (Fig. 
176).  Venter  black  surrounded  by  yellow. 
Wide  white  median  line  with  pair  of  large 
white  spots  on  either  side  of  spiracle. 
Sometimes  with  parallel  pair  of  lateral 
white  lines  that  join  transverse  white  line 
posteriorly  (Fig.  177).  Sternum  black  with 
wide,  white  line  widening  anteriorly,  often 


broken  in  center  (Fig.  177).  Ratio  of  eye 
diameters:  posterior  medians  and  anterior 
medians  0.9,  anterior  laterals  1.3,  posterior 
laterals  1.2.  Anterior  median  eyes  separat- 
ed by  1.2  diameters,  posterior  median  eyes 
by  0.6,  anterior  median  eyes  separated 
from  anterior  laterals  by  3.4  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.4  their  diameters.  Total  length  8.3 
mm.  Carapace  4  mm  long,  3.3  wide.  First 
femur  4.5  mm,  patella  and  tibia  4.8,  meta- 
tarsus 4.5,  tarsus  1.3.  Second  patella  and 
tibia  4.3  mm,  third  2.5,  fourth  3.8. 

Male  from  Baja  California  Norte,  Mex- 
ico. Male  carapace,  dorsum,  venter,  ster- 
num as  in  female  (Figs.  174,  175).  Femur 
I  with  row  of  five  macrosetae  on  anterior 
side;  row  of  nine  macrosetae  on  anterov- 
entral side.  Ratio  of  eye  diameters:  poste- 
rior medians  and  anterior  medians  0.8,  an- 
terior laterals  1.1,  posterior  laterals  1.3. 
Anterior  median  eyes  separated  by  1  di- 
ameter, posterior  median  eyes  by  0.8,  an- 
terior median  eyes  separated  from  anterior 
laterals  by  2.1  diameters  of  anterior  lateral 
eyes,  lateral  eyes  separated  by  0.2  their  di- 
ameters. Total  length  5.5  mm.  Carapace 
2.8  mm  long,  2.3  wide.  First  femur  4.5 
mm,  patella  and  tibia  4.5,  metatarsus  4.5, 
tarsus  1.4.  Second  patella  and  tibia  3.8 
mm,  third  1.9,  fourth  3. 

Diagnosis.  Within  the  M.  incrassata  spe- 
cies group,  only  M.  gosoga,  M.  inca,  and 
M.  maya  have  projecting  distal  embolic 
apophyses  (Figs.  157,  164,  171).  Of  these, 
only  M.  gosoga  has  a  distinctly  pointed  one 
(Fig.  171).  The  epigynum  of  M.  gosoga  dif- 
fers from  those  of  other  species  in  the  M. 
incrassata  group  by  the  small,  oval  shape 
of  the  openings  (Fig.  173). 

Variation.  Average  body  length  of  two 
females  examined  6.8  mm,  range  5  to  8.5 
mm.  Average  body  length  of  two  males  ex- 
amined 4.9  mm,  range  4.1  to  5.7  mm. 

Natural  History.  Mature  specimens 
have  been  collected  in  June  through  Au- 
gust (Fig.  322)  between  500  and  2,000  m. 
Levi  (1977)  notes  that  these  have  been 
collected  on  Opuntia  and  desert  vegeta- 
tion. 


60 


Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


Distribution.  Southwestern  U.S.  and 
Baja  California  Norte,  Mexico  (Levi,  1977, 
map  1;  Map  9). 

Records  Examined.  MEXICO  BaJa  Calif.  Norte:  23 
mi  S  Catavina,  29°32'N,  114°57'W,  20.vi.l983  (L. 
Strange  &  R.  Miller,  FSCA).  USA  Arizona:  Show 
Low  Lake,  34°12'N,  110°0'W,  15.viii.l966  (MCZ). 

22.  Metepeira  olmec  new  species 
Figures  178-184,  324;  Map  11 

Holotype.  Female  from  Foitin  de  las  Flores,  Vera- 
cmz,  Mexico,  26.vi.1944,  L.  L  Davis,  in  AMNH. 
The  specific  name  is  a  noun  in  apposition  after  the 
ancient  Indian  people  of  southern  Veracruz  and  Ta- 
basco. 

Description.  Female  holotype.  Carapace 
reddish  brown  with  white  region  around 
eyes  that  extends  posteriorly  along  lateral 
margins  (Fig.  183).  Legs  ringed  on  distal 
ends  of  articles.  Femur  I  with  row  of  four 
macrosetae  on  anterior  side;  none  on  an- 
teroventral  side.  Dorsum  of  abdomen  with 
usual  Metepeira  folium  (Fig.  183).  Venter 
of  abdomen  with  wide,  long  white  median 
line  set  within  surrounding  U-shaped 
markings.  Pair  of  white  spots  on  either 
side  of  spiracle  (Fig.  184).  Sternum  has 
wide  median  white  line  widening  anteri- 
orly with  broken  constriction  in  center 
(Fig.  184).  Ratio  of  eye  diameters:  poste- 
rior medians  and  anterior  medians  1.0,  an- 
terior laterals  1.1,  posterior  laterals  1.5. 
Anterior  median  eyes  separated  by  1.5  di- 
ameters, posterior  median  eyes  by  0.8,  an- 
terior median  eyes  separated  from  anterior 
laterals  by  2.1  diameters  of  anterior  lateral 
eyes,  lateral  eyes  separated  by  0.2  their  di- 
ameters. Total  length  5.9  mm.  Carapace 
2.8  mm  long,  2  wide.  First  femur  2.8  mm, 
patella  and  tibia  2.8,  metatarsus  2.5,  tarsus 
1.  Second  patella  and  tibia  2.5  mm,  third 
1.5,  fourth  2.3. 

Male  paratype  from  Fortin  de  las  Flo- 
res,  Veracruz,  Mexico.  Carapace  dark  with 
light  region  around  eyes  and  a  light  mark 
anterior  to  thoracic  furrow  (Fig.  181). 
Legs  darker  than  in  female.  Macrosetae  on 
legs  variable  with  spiders  size — femur  I 
with  row  of  three  or  four  macrosetae  on 
anterior  side;  one  to  three  on  anteroven- 


tral  side.  Dorsum,  venter,  sternum  as  in 
female,  except  median  white  line  on  ven- 
ter shortened  to  spot  and  median  white 
line  on  sternum  often  broken  (Figs.  181, 
182).  Ratio  of  eye  diameters:  posterior  me- 
dians and  anterior  medians  1.0,  anterior 
laterals  1.3,  posterior  laterals  1.3.  Anterior 
median  eyes  separated  by  1.3  diameters, 
posterior  median  eyes  by  0.8,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  1.3  diameters  of  anterior  lateral  eyes, 
lateral  eyes  separated  by  0.1  their  diame- 
ters. Total  length  2.8  mm.  Carapace  1.4 
mm  long,  1.2  wide.  First  femur  1.7  mm, 
patella  and  tibia  1.6,  metatarsus  1.1,  tarsus 
0.7.  Second  patella  and  tibia  1.3  mm,  third 
0.8,  fourth  1.1. 

Diagnosis.  Female  M.  olmec  are  diag- 
nosed by  their  almost  perfectly  round  epi- 
gynal  depressions  with  uniform  thickness 
around  the  posterior  edges  (Fig.  180).  The 
male  embolus  is  relatively  short  and  has  a 
vestigial  distal  embolic  apophysis  that  does 
not  project  foi^ward,  but  instead  drops  off, 
forming  a  shai^  cuive  (Fig.  178).  Among 
other  species  in  the  M.  incrassata  species 
group,  this  embolus  shape  is  also  seen  in 
M.  conianche  and  M.  triangularis:  unlike 
M.  conianche  (Fig.  185),  the  darker,  scler- 
otized  portion  of  the  embolus  does  not  ex-\ 
tend  over  the  hump  of  the  distal  embolic 
apophysis;  unlike  M.  triangularis  (Fig. 
199),  the  flagellae  are  thicker  than  the 
base  of  the  median  apophysis. 

Variation.  Average  body  length  of  four 
females  examined  6.2  mm,  range  5.6  to  6.7 
mm.  Average  body  length  of  three  males 
examined  2.3  mm,  range  2.2  to  2.6  mm. 

Natural  History.  Mature  specimens 
have  been  collected  perennially  (Fig.  324) 
at  elevations  between  500  and  1,400  m. 

Distribution.  Montane  rain  forests  from 
Veracruz  to  Panama  (Map  11). 

Records  Examined.  COSTA  RICA  San  Jose:  San 
Antonio  de  Escanza,  9°59'N,  84°11'W  [?],  4.iii.l984 
(W.  Eberhard,  MCZ).  MEXICO  Veracruz:  Foitin  de 
las  Flores,  18°54'N,  97°0'W,  26.vi.1944  (L.  I.  Davis, 
AMNH);  Papantla,  20°27'N,  97°19'W,  12.x.  1947  (H. 
Wagner,  AMNH).  PANAMA  Chiriqut:  Boquete, 
8°47'N,    82°26'W,    15.viii.l950    (A.    M.    Chickering, 


Metepeira  •  Piel         61 


gosoga 
(21) 


184 

olmec 

(22) 


190  ^  191 

Comanche 
(23) 


Figures  171-177.  Metepeira  gosoga  CnamberWn  and  Ivie  (sp.  21;  29°32'N,  114°57'W).  171,  male  palpus,  mesal.  172,  epigynum, 

posterior.  173,  epigynum,  ventral.  174,  male,  dorsal.  175,  male,  ventral.  176,  female,  dorsal.  177,  female,  ventral. 

Figures  178-184.  Metepeira  olmec  new  species  (sp.  22;  18°54'N,  97°0'W).  178,  male  palpus,  mesal.  179,  epigynum,  posterior. 

180,  epigynum,  ventral.  181,  male,  dorsal.  182,  male,  ventral.  183,  female,  dorsal.  184,  female,  ventral. 

Figures  185-191.  Metepeira  comancfie  Levi  (sp.  23  [185,188,189]  33°22'N,  99°56'W;  [186,187]  25°45'N,  10r55'W;  [190, 191] 

26°41'N,  101°23'W).  185,  male  palpus,  mesal.  186,  epigynum,  posterior.  187,  epigynum,  ventral.  188,  male,  dorsal.  189,  male, 

ventral.  190,  female,  dorsal.  191,  female,  ventral. 

Scale  bars:  dorsum  and  venter  figures  1 .0  mm. 


62         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


MCZ),  7.viii.l954  (A.  M.  Chickering,  MCZ).  Panama: 
Cerro  Galera,  8°55'N,  79°38'W,  7.i.l977  (H.  W.  Levi 
&  Y.  Lubin,  MCZ). 

23.  Metepeira  comanche  Levi 
Figures  185-191,  331;  IVlap  11 

Metepeira  comanche  Levi,  1977:  204,  figs.  61-69,  S, 
? .  Male  holotype  from  9.7  km  west  of  O'Brien, 
Haskell  Co.,  Texas,  in  the  MCZ,  examined.  Brig- 
noli,  1983:  275. 

Description.  Female  from  Gloria,  Coa- 
huila,  Mexico.  Carapace  dark  with  light  re- 
gion around  eyes,  extending  posteriorly 
behind  posterior  lateral  eyes  (Fig.  190). 
Legs  ringed.  Femur  I  with  row  of  four  ma- 
crosetae  on  anterior  side;  one  or  two  on 
anteroventral  side.  Dorsal  folium  high  in 
contrast;  black  comma-shaped  markings 
shadow  white  fleur-de-lis  pattern  (Fig. 
190).  Venter  of  abdomen  with  wide  white 
median  line  posteriorly  set  within  sur- 
rounding U-shaped  marking.  Pair  of  white 
spots  on  either  side  of  spiracle  (Fig.  191). 
Sternum  has  wide  median  white  line  wid- 
ening anteriorly  (Fig.  191).  Ratio  of  eye 
diameters:  posterior  inedians  and  anterior 
medians  0.9,  anterior  laterals  1.2,  posterior 
laterals  1.1.  Anterior  median  eyes  separat- 
ed by  1.2  diameters,  posterior  median  eyes 
by  1.0,  anterior  median  eyes  separated 
from  anterior  laterals  by  2.7  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.2  their  diameters.  Total  length  7.8 
mm.  Carapace  3.3  mm  long,  2.5  wide. 
First  femur  3.6  mm,  patella  and  tibia  3.9, 
metatarsus  3.4,  tarsus  1.2.  Second  patella 
and  tibia  3.3  mm,  third  2,  fourth  2.9. 

Male  from  9.7  km  west  of  O'Brien,  Has- 
kell Co.,  Texas.  Carapace  light  around 
eyes,  darker  posteriorly  (Fig.  188).  Legs 
ringed.  Macrosetae  on  femur  I  variable; 
usually  row  of  four  macrosetae  on  anterior 
side,  six  on  anteroventral  side.  Dorsum 
with  usual  Metepeira  folium  (Fig.  188). 
Venter  with  median  oval  white  mark;  pair 
of  white  spots  on  either  side  of  spiracle 
(Fig.  189).  Median  white  line  on  sternum 
usually  broken  (Fig.  189).  Ratio  of  eye  di- 
ameters: posterior  medians  and  anterior 
medians  1.0,  anterior  laterals  1.3,  posterior 


laterals  1.3.  Anterior  median  eyes  separat- 
ed by  1.4  diameters,  posterior  median  eyes 
by  0.9,  anterior  median  eyes  separated 
from  anterior  laterals  by  1.6  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.2  their  diameters.  Total  length  3.1 
mm.  Carapace  1.7  mm  long,  1.3  wide. 
First  femur  2.4  mm,  patella  and  tibia  2.2, 
metatarsus  2,  tarsus  0.9.  Second  patella 
and  tibia  1.9  mm,  third  1.0,  fourth  1.5. 

Diagnosis.  The  epigynum  of  M.  com- 
anche resembles  those  of  M.  may  a  and  M. 
inca  because  the  oval  epigynal  openings 
are  formed  out  of  membranous  surfaces 
that  are  distinctly  Separate  from  the  base 
of  the  scape  (Figs.  159,  166,  187).  The  epi- 
gynum of  M.  comanche  differs  from  M. 
niaya  and  M.  inca  by  having  much  narrow- 
er epigynal  openings  (compare  Fig.  187 
with  Figs.  159,  166).  The  male  embolus  is 
relatively  short  and  has  a  vestigial  distal 
embolic  apophysis  that  does  not  project 
forward,  but  instead  drops  off,  forming  a 
sharp  curve  (Fig.  185).  Among  other  spe- 
cies in  the  M.  incrassata  species  group, 
this  embolus  shape  is  also  seen  in  M.  olmec 
and  M.  triangularis:  unlike  M.  olmec  (Fig. 
178),  the  darker,  sclerotized  portion  of  the 
embolus  extends  over  the  hump  of  the  dis- 
tal embolic  apophysis;  unlike  M.  triangu- 
laris (Fig.  199),  the  flagellae  and  base  of 
the  median  apophysis  are  wide  (Fig.  185). 

Natural  Histortj.  Mature  specimens 
have  been  collected  from  May  to  Novem- 
ber (Levi,  1977,  Fig.  331). 

Distribution.  New  Mexico,  Texas,  and 
northern  Mexico  (Levi,  1977,  map  1;  Map 
11). 

Records  Examined.  MEXICO  Coahuila:  Gloria, 
26°41'N,  101°23'W,  24.viii.1947  (W.  J.  Gerisch, 
AMNH);  Paila,  25°45'N,  101°55'W,  21.viii.l947  (W. 
J.  Gertsch,  AMNH). 

24.  Metepeira  pimungan  new  species 
Figures  192-198,  330;  Map  9 

Holotype.  Male  from  San  Miguel  Island,  Santa  Bar- 
bara Co.,  California,  USA,  20.vii.l968,  M.  E. 
Thompson,  in  the  MCZ.  The  specific  name  is  a 
noun  in  apposition  after  the  Indian  people  who 
once  lived  on  the  Channel  Islands. 


Metepeira  •  Piel 


63 


Description.  Female  paratype  from  San 
Miguel  Island,  California  USA.  Carapace 
with  large  white  eye  region  and  lateral 
posterior  extensions.  White  median  arrow- 
shaped  mark  extends  to  thoracic  furrow 
(Fig.  197).  Legs  mostly  white  with  rings  on 
legs  III  and  IV.  Femur  I  with  row  of  four 
macrosetae  on  anterior  side;  one  on  anter- 
oventral  side.  Dorsum  of  abdomen  with 
usual  Metepeira  folium  (Fig.  197);  venter 
wide,  with  long  white  median  line,  flanked 
by  shorter  parallel  thin  white  lines  on  ei- 
ther side.  Pair  of  white  spots  on  either  side 
of  spiracle.  Sternum  black  with  posterior 
drop-shaped  white  mark  (Fig.  198).  Ratio 
of  eye  diameters:  posterior  medians  and 
anterior  medians  1.0,  anterior  laterals  1.4, 
posterior  laterals  1.3.  Anterior  median 
eyes  separated  by  1.6  diameters,  posterior 
median  eyes  by  1.0,  anterior  median  eyes 
separated  from  anterior  laterals  by  3.4  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.3  their  diameters.  To- 
tal length  6.9  mm.  Carapace  3.3  mm  long, 
2.6  wide.  First  femur  3.5  mm,  patella  and 
tibia  3.5,  metatarsus  2.8,  tarsus  1.3.  Sec- 
ond patella  and  tibia  3  mm,  third  1.9, 
fourth  2.8. 

Male  holotype.  Carapace,  legs,  abdo- 
men, venter,  sternum  as  in  female,  though 
legs  darker  and  not  ringed  (Figs.  195, 
196).  Femur  I  with  row  of  four  to  five  ma- 
crosetae on  anterior  side;  four  to  five  on 
antero ventral  side:  both  rows  concentrated 
distally  on  femur.  Ratio  of  eye  diameters: 
posterior  medians  and  anterior  medians 
1.0,  anterior  laterals  1.3,  posterior  laterals 
1.3.  Anterior  median  eyes  separated  by  1.6 
diameters,  posterior  median  eyes  by  0.8, 
anterior  median  eyes  separated  froin  an- 
terior laterals  by  1.9  diameters  of  anterior 
lateral  eyes,  lateral  eyes  separated  by  0.3 
their  diameters.  Total  length  4.5  mm.  Car- 
apace 2.3  mm  long,  1.8  wide.  First  femur 
3.1  mm,  patella  and  tibia  3,  metatarsus  2.8, 
tarsus  1.  Second  patella  and  tibia  2.5  mm, 
third  1.3,  fourth  1.9. 

Diagnosis.  Like  M.  atascadero,  but  un- 
like other  species  in  the  M.  incrassata 
group,  the  posterior  end  of  the  sternum  of 


M.  pimungan  has  a  white  water  drop  mark 
(Figs.  196,  198).  Unlike  M.  atascadero,  M. 
pimungan  lacks  U-shaped  white  markings 
on  the  venter,  just  anterior  to  the  spiracle 
(compare  Fig.  198  with  Fig.  221).  In  con- 
trast to  other  species  in  the  M.  incrassata 
group,  the  embolus  on  male  M.  pimungan 
speciinens  is  curved  to  forin  a  gentle  S- 
shape  (Fig.  192).  The  female  has  a  squar- 
ish-shaped black  mark  inside  each  epigyn- 
al  depression  (Fig.  198)  which  is  not  seen 
in  other  M.  incrassata  group  species. 

Variation.  Average  body  length  of  four 
females  examined  7.7  mm,  range  6.9  to  8.5 
mm.  Average  body  length  of  five  inales  ex- 
amined 5.3  mm,  range  4.6  to  6.4  mm. 

Natural  History.  Mature  specimens 
have  been  collected  in  August  (Fig.  330). 
Webs  are  found  near  the  ground,  shel- 
tered from  the  perennial  strong  winds  by 
Cortjopsis  and  lupines. 

Distribution.  Endemic  to  San  Miguel 
Island  (Map  9). 

Records  Examined.  USA  California:  San  Miguel  Is- 
land, 34°2'28.6"N,  120°21'13.6"W,  ll.viii.l995"(W.  H. 
Piel,  MCZ);  34°2'6.1"N,  120°21'9.1"W,  12.viii.l995 
(W.  H.  Piel,  MCZ);  34°2'N,  120°22'W,  20.viii.l968 
(M.  E.  Thompson,  MCZ);  34°3'3.1"N,  120°21'53.3"W, 
13.viii.l995  (W.  H.  Piel,  MCZ);  behind  dunes  on 
beach,  34°2'43.8"N,  120°21'0.8"W,  ll.viii.l995  (W.  H. 
Piel,  MCZ). 

25.  Metepeira  triangularis  (Franganillo) 
Figures  199-205,  337;  Map  14 

Mangora  triangularis  Franganillo,  1930:  21—22.  Fe- 
male holotype  from  Sierra  Maestra,  lost.  Platnick, 
1993:  449. 

Metepeira  labijrinthea: — Franganillo,  1936:  75.  Bry- 
ant, 1940:  341.  Platnick,  1993:  449. 

Metepeira  triangidaris: — Archer,  1958:  15,  fig.  37,  9 . 

Metepeira  acostai  Archer,  1958:  15,  fig.  36,  9 .  Female 
holotype  from  the  savannas,  Agramont,  Camagiiey 
Province,  Cuba,  in  the  AMNH,  examined.  NEW 
SYNONYMY. 

Metepeira  hani  Archer,  1965:  132,  figs.  11,  17  9,  13 
S .  Male  and  female  syntypes  from  Bani,  Domini- 
can Republic,  in  the  AMNH,  examined.  NEW 
SYNONYMY.  Brignofi,  1983:  275. 

Note.  Franganillo's  collection  vials  are  not  la- 
beled, making  it  impossible  to  find  the  holotype. 
Based  on  the  descriptions  by  Franganillo  (1930), 
Archer  (1958),  the  illustrations  of  Archer  (1958, 
1965),  and  material  determined  by  Archer,  I  con- 
clude that  there  is  no  evidence  to  support  three 


64         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


separate  species.  For  the  name  M.  triangularis,  I 
designate  a  female  neotype  from  Camagiiey  Prov- 
ince, Cuba.  21°30'N,  78°10'W,  x.l954  (J.  T.  Acosta, 

AMNH). 

Description.  Female  from  La  Descu- 
bierta,  Lago  Enriquillo,  Isla  Cabritos,  Do- 
minican Republic.  Light  reddish  brown 
carapace,  white  around  eyes  (Fig.  204). 
Legs  yellowish  white,  sometimes  ringed 
brown  at  distal  ends  of  articles.  Femur  I 
with  row  of  three,  or  sometimes  four,  ma- 
crosetae  on  anterior  side;  none  on  anter- 
oventral  side.  Dorsal  folium  white  with 
slightly  darker  markings  posteriorly  (Fig. 
204).  In  some  cases,  dark  anterior  marks 
on  shoulders  of  dorsum.  Venter  of  abdo- 
men black  with  wide  white  median  line, 
sometimes  flanked  by  thinner  white  lines 
that  together  form  an  anchor  shape  (Fig. 
205).  Pair  of  small  white  spots  on  either 
side  of  spiracle.  Sternum  black  with  me- 
dian white  line,  wider  anteriorly  (Fig.  205). 
Ratio  of  eye  diameters:  posterior  medians 
and  anterior  medians  1.0,  anterior  laterals 
1.4,  posterior  laterals  1.4.  Anterior  median 
eyes  separated  by  1.5  diameters,  posterior 
median  eyes  by  1.1,  anterior  median  eyes 
separated  from  anterior  laterals  by  3.3  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.2  their  diameters.  To- 
tal length  5.9  mm.  Carapace  2.7  mm  long, 
2  wide.  First  femur  2.6  mm,  patella  and 
tibia  2.9,  metatarsus  2.5,  tarsus  1.  Second 
patella  and  tibia  2.4  mm,  third  1.4,  fourth 
2.2. 

Male  from  La  Descubierta,  Lago  Enri- 
quillo, Isla  Cabritos,  Dominican  Republic. 
Carapace  as  in  female,  though  lighter  eye 
region  often  extends  to  cover  entire  ante- 
rior half  (Fig.  202).  Legs  yellowish  white. 
Distal  half  of  femora  often  dark.  Femur  I 
with  row  of  four  macrosetae  on  anterior 
side;  three  to  five  on  anteroventral  side. 
Dorsal  folium  white  witli  slightly  darker 
markings  posteriorly  (Fig.  202).  Venter  of 
abdomen  as  in  female;  sternum  black  with 
median  white  line  (Fig.  203).  Ratio  of  eye 
diameters:  posterior  medians  and  anterior 
medians  1.1,  anterior  laterals  0.7,  posterior 
laterals  1.1.  Anterior  median  eyes  separat- 


ed by  1.6  diameters,  posterior  median  eyes 
by  0.8,  anterior  median  eyes  separated 
from  anterior  laterals  by  1.0  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.1  their  diameters.  Total  length  4.2 
mm.  Carapace  2  mm  long,  1.5  wide.  First 
femur  2.9  mm,  patella  and  tibia  3,  meta- 
tarsus 2.8,  tarsus  1.1.  Second  patella  and 
tibia  2.4  mm,  third  0.8,  fourth  1.3. 

Diagnosis.  The  epigynum  of  M.  trian- 
gularis differs  from  other  species  by  the 
thin,  sclerotized  openings  in  the  shape  of 
Ray-Ban  sunglasses  (Fig.  201).  The  male 
embolus  has  a  vestigial  distal  embolic 
apophysis  that  does  not  project  forward, 
but  instead  drops  off,  forming  a  sharp 
cui-ve  (Fig.  199).  Among  other  species  in 
the  M.  incrassata  species  group,  shorter 
versions  of  this  embolus  shape  is  also  seen 
in  M.  Comanche  and  M.  olmec:  unlike  M. 
Comanche  (Fig.  185),  the  darker,  sclero- 
tized portion  of  the  embolus  does  not  ex- 
tend over  the  hump  of  the  distal  embolic 
apophysis;  unlike  M.  olmec  (Fig.  178),  the 
flagellae  and  base  of  the  median  apophysis 
are  much  thinner  (Fig.  199). 

Variation.  Average  body  length  of  seven 
females  examined  5.5  mm,  range  4.6  to  6.3 
mm.  Average  body  length  of  five  males  ex- 
amined 3.8  mm,  range  3.2  to  4.3  mm.       j 

Natural  History.  Mature  specimens 
have  been  collected  from  June  to  Febru- 
ary, although  I  suspect  that  like  other  Ca- 
ribbean species,  M.  triangularis  is  actually 
entirely  perennial  (Fig.  337). 

Distribution.  Dominican  Republic  and 
Cuba  (Map  14). 

Records  Examined.  CUBA  Camagiiey  Prov.: 
21°30'N,  78°10'W,  X.1954  (J.  T.  Acosta,  AMNH).  Ma- 
tanzas:  Matanzas,  Parque  Watldns,  23°3'N,  81°35'W, 
9.viii.l955  (A.  F.  Archer,  AMNH);  Pan  de  Palenque 
[?],  23°1'N,  81°43'W,  ll.viii.l955  (A.  F.  Archer, 
AMNH).  Oriente:  Banes,  20°58'N,  75°43'W, 
2.viii.l955  (A.  F  Archer,  AMNH);  Cuabitas,  Santiago, 
20°4'N,  75°48'W,  15.xii.l955  (P  Alayo,  AMNH); 
Puerto  Boniato,  20°7'N,  75°47'W,  4.xi.l945  (P  Alayo, 
AMNH).  Sierra  las  Casas:  Isla  de  Pinos,  21°53'N, 
82°48'W,  17.viii.l955  (A.  F  Archer,  AMNH).  DO- 
MINICAN REPUBLIC  Azua:  Hatillo,  18°24'N, 
70°32'W,  30.i.l991  (FelLx  E.  Del  Monte,  CAS).  Ban- 
ahona:  Sierra  Martin  Garcia  [?],  18°25'N,  70°30'W, 
8.viii.l958    (A.    F    Archer   &    E.    de    Boyrie    Moya, 


Metepeira  •  Piel 


65 


pimungan 
(24) 


A. 

9 


triangularis 

(25) 


212 

arizonica 
(26) 

Figures  192-198.  Metepeira  pimungan  new  species  (sp.  24;  34°2'N,  120°22'W).  192,  male  palpus,  mesal.  193,  epigynum, 

posterior.  194,  epigynum,  ventral.  195,  male,  dorsal.  196,  male,  ventral.  197,  female,  dorsal.  198,  female,  ventral. 

Figures  199-205.  Metepeira  triangularis  (Franganillo)  (sp.  25;  18°34'N,  71°44'W).  199,  male  palpus,  mesal.  200,  epigynum, 

posterior.  201,  epigynum,  ventral.  202,  male,  dorsal.  203,  male,  ventral.  204,  female,  dorsal.  205,  female,  ventral. 

Figures  206-212.  Metepeira  arizonica  Chamberlin  and  Ivie  (sp.  26;  28°53'N,  1 13°4'W).  206,  male  palpus,  mesal.  207,  epigynum, 

posterior.  208,  epigynum,  ventral.  209,  male,  dorsal.  210,  male,  ventral.  211,  female,  dorsal.  212,  female,  ventral. 

Scale  bars:  dorsum  and  venter  figures  1 .0  mm. 


66 


Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


AMNH).  La  Independencia:  2  km  SE  El  Limon,  Ji- 
mani.  S  side  of  road,  18°25'N,  71°45'W,  30.i.l991 
(Felix  E.  Del  Monte,  MNSD);  Isla  Cabritos:  La  Des- 
cubierta,  Lago  Enriquillo,  18°34'N,  71°44'W, 
18.vi.l981  (E.  Marcano,  MNSD).  Montecristi:  Car- 
retera  Montecristi,  El  Morro  [?],  19°54'N,  71°39'W, 
2.xii.l991  (EelLx  E.  Del  Monte,  MNSD),  21.xii.l991 
(FelLx  E.  Del  Monte,  MNSD).  Peravia:  Las  Dunas, 
Prov.  Bani  [?],  18°25'N,  71°25'W,  23.i.l992  (Felix  E. 
Del  Monte,  MNSD).  Frov.  Tnijillo  Valdez:  W  of 
Bani,  18°17'N,  70°22'W,  8.viii.l958  (A.  F.  Archer, 
AMNH).  San  Juan:  1  km  S  Las  Matas  de  Farfan, 
18°51'N,  71°31'W,  25.viii.1970  (B.  Patterson,  MCZ). 

26.  Metepeira  arizonica 
Chamberlin  and  Ivie 
Figures  206-213,  319;  Map  9 

Metepeira  arizonica  Chamberlin  and  Ivie,  1942:  69, 
figs.  182-187,  9,6.  Female  holotype  from  Canyon 
Lake,  Arizona,  in  the  AMNH,  examined.  Levi, 
1977:  200,  figs.  12,  13,  39^6,  9,3.  Brignoh,  1983: 
275. 

Description.  Female  from  Isla  Partida, 
Baja  Calif.  Norte,  Mexico.  General  color- 
ation high  in  contrast.  Carapace  dark 
brown;  large  white  eye  region,  white  lat- 
eral posterior  extensions,  and  short  median 
posterior  extension  (Fig.  211).  Distinct 
black  rings  on  all  legs.  Femur  I  with  row 
of  four  macrosetae  on  anterior  side;  four 
on  anteroventral  side.  Dorsum  of  abdo- 
men with  usual  Metepeira  folium,  though 
generally  slightly  lighter  (Fig.  211);  venter 
wide,  with  long  white  median  line  set 
within  surrounding  U-shaped  markings. 
Pair  of  white  spots  on  either  side  of  spi- 
racle (Fig.  212).  Wide  median  white  line 
widening  anteriorly  with  constriction  in 
center,  set  on  black  sternum  (Fig.  212). 
Ratio  of  eye  diameters:  posterior  medians 
and  anterior  medians  1.0,  anterior  laterals 
1.4,  posterior  laterals  1.2.  Anterior  median 
eyes  separated  by  1.4  diameters,  posterior 
median  eyes  by  0.9,  anterior  median  eyes 
separated  from  anterior  laterals  by  2.7  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.1  their  diameters.  To- 
tal length  6.2  mm.  Carapace  2.8  mm  long, 
2.2  wide.  First  femur  3.4  mm,  patella  and 
tibia  3.5,  metatarsus  3,  tarsus  1.2.  Second 
patella  and  tibia  3  mm,  third  1.9,  fourth 
2.8. 


Male  from  Isla  Partida,  Baja  Calif.  Nor- 
te, Mexico.  Carapace,  legs,  abdoiTien,  ven- 
ter, sternum  as  in  female,  though  legs 
darker  with  less  distinct  rings,  and  white 
line  on  sternum  often  broken  (Figs.  209, 
210).  Femur  I  with  row  of  four  macrosetae 
on  anterior  side;  five  to  eight  on  anterov- 
entral side.  Ratio  of  eye  diameters:  poste- 
rior medians  and  anterior  medians  1.1,  an- 
terior laterals  1.4,  posterior  laterals  1.2. 
Anterior  median  eyes  separated  by  1.5  di- 
ameters, posterior  median  eyes  by  0.7,  an- 
terior median  eyes  separated  from  anterior 
laterals  by  1.6  diameters  of  anterior  lateral 
eyes,  lateral  eyes  separated  by  0.2  their  di- 
ameters. Total  length  4.4  mm.  Carapace 
2.3  mm  long,  1.8  wide.  First  femur  3.3 
mm,  patella  and  tibia  3.3,  metatarsus  3.2, 
tarsus  1.2.  Second  patella  and  tibia  2.8 
mm,  third  1.5,  fourth  2.2. 

Diagnosis.  Unlike  all  other  species  in 
the  M.  incrassata  group,  the  epigynum  of 
M.  arizonica  is  puffy  and  swollen,  causing 
the  openings  to  reduce  (from  a  ventral 
view)  to  crescent-shaped  slits  (Fig.  187). 
The  embolus  differs  from  others  in  the  M. 
incrassata  group  because  the  distal  em- 
bolic apophysis  is  reduced  to  a  gentle  arch- 
ing cui-ve  (Fig.  206). 

Variation.  Average  body  length  of  nine 
females  examined  5.5  mm,  range  4.7  to  6.3 
mm.  Average  body  length  of  13  males  ex- 
amined 3.9  mm,  range  3.1  to  5.1  mm.  In 
some  males  a  small  ventral  keel  is  visible 
extending  beyond  the  flagella  of  the  me- 
dian apophysis  (Levi,  1977,  fig.  46). 

'Natural  History.  Mature  specimens  are 
most  commonly  collected  between  April 
and  August  (Fig.  319),  from  dry  oak-pine- 
juniper  woodland,  alfalfa,  cactus,  and  de- 
sert shrub  (Levi,  1977).  Localities  have  a 
large  range  in  altitude,  from  sea  level  to 
2,300  m;  however,  spiders  from  southern 
regions  live  at  higher  elevations  than  spi- 
ders from  northern  regions  (Fig.  213). 

Distribution.  American  southwest  to 
San  Luis  Potosi  (Levi,  1977,  map  1;  Map 
9). 

Records  Examined.  MEXICO  Baja  Calif.  Norte:  15 


Metepeira  •  Piel 


67 


o 

o 

1 

2200  • 

^^ 

° 

B 

1800  - 

° 

^^^ 

o 

o 
1 

1400  ■ 
1000  • 

"""V 

»^ 

i 

■^^ 

° 

1 

600- 
200- 

— ma — a__- 

nn.  a.  0  .  . . . — ^ 

Latitude  (degrees  north) 

Figure  213.  The  elevation  of  collection  localities  for  M.  arizon- 
ica  at  their  corresponding  latitudes.  Species-specific  altitudes 
appear  to  decrease  with  distance  from  the  equator.  Elevations 
estimated  from  NOAA  database  of  5-  by  5-minute  geographic 
tiles. 


mi  S  Punta  Prieta,  28°58'N,  114°17'W,  7.vii.l973  (S. 
C.  Williams  &  K.  B.  Blair,  CAS);  24  mi  S  Santa  Ines, 
29°20'N,  114°20'W,  7.vii.l973  (S.  C.  Williams  &  K. 

B.  Blair,  CAS);  26  mi  S  San  Felipe,  30°38'N, 
114°50'W,  15.iv.l965  (D.  Q.  Cavagnaro,  C.  E.  &  E. 
S.  Ross,  V.  L.  Vesterby,  CAS);  6  km  NW  Racho  Santa 
Ines,  29°43'N,  114°43'W,  28.iii.1981  (Paul  E.  Blom, 
WHO;  6  mi  S  San  Felipe,  30°55'N,  114°52'W 
14.iv.l965  (D.  Q.  Cavagnaro,  C.  E.  &  E.  S.  Ross,  V. 
L.  Vesterby,  CAS);  9  km  NW  Racho  Santa  Ines, 
29°46'N,  114°46'W,  3.vii.l981  (David  Crowe,  WHC); 
Arroyo  de  Calamajue  at  carbonated  spring,  29°38'N, 
114°25'W,  26.vi.1973  (S.  C.  Williams  &  K.  B.  Blair, 
CAS);  Canon  de  Cuadalupe,  off  Laguna  Salada, 
32°10'N,  115°48'W,  ll.i.l958  (V.  Roth,  AMNH);  El 
Mayor,  32°5'N,  115°13'W,  15.vi.l952  (M.  Cazier,  W 
Gertsch,  &  R.  Schrammel,  AMNH);  Isla  Mejia, 
29°34'N,  113°35'W,  30.ivl921  (J.  C.  Chamberlin, 
MCZ);  Isla  Partida,  28°53'N,  113°4'W,  2.vii.l921  (J. 

C.  Chamberlin,  MCZ);  Isla  San  Lorenzo,  N  end, 
28°40'N,  112°52'W,  24.vi.1921  (J.  C.  Chamberlin, 
CAS);  Isla  San  Lorenzo,  south  Tobart,  28°36'N, 
112°46'W  [?],  9.iv.l921  (J.  C.  Chamberlin,  CAS); 
Puerto  Refugio,  N  end  Angel  de  la  Guarda  Isl.  Webs 
in  cracks  in  cliff  overhanging  beach,  29°34'N, 
113°32'W,  18.iv.l962  (Howard  W.  Campbell,  CAS); 
San  Pedro  Martir,  30°45'N,  115°13'W,  18.iv.l921  (J. 
C.  Chamberlin,  CAS).  Coahuila:  25  mi  SE  San  Pedro, 
25°35'N,  102°50'W,  21.viii.l947  (W.  J.  Gertsch, 
AMNH);  Cabos,  25°35'N,  101°43'W,  21.viii.l947  (W. 
J.  Gertsch,  AMNH);  Gloria,  26°41'N,  10r23'W, 
24.viii.1947  (W  J.  Gertsch,  AMNH);  Saltillo, 
25°25'N,  101°0'W,  23.V.1952  (M.  Cazier,  W  Gertsch, 
&  R.  Schrammel,  AMNH).  Durango:  La  Loma, 
25°27'N,  103°40'W,  20.viii.l947  (W  J.  Gertsch, 
AMNH);  Providencia,  26°44'N,  105°56'W 
24.viii.1947  (A.  M.  Davis,  AMNH);  San  Isidro,  60  mi 
NW  Durango,  25°1'N,  105°6'W,  19.viii.l947  (W  J. 
Gertsch,  AMNH).  San  Luis  Potosi:  3  km  W  Pilares, 
21°55'34"N,  100°48'6"W,  21.X.1994  (W  H.  Piel, 
MCZ).  Sinaloa:  Las  Saleras  Is.  (=  Isla  SaHaca  [?]), 
25°11'N,  108°20'W  [?],  13.vi.l921  (J.  C.  Chamberlin, 


CAS).  Sonora:  20  mi  N  Hermosillo,  29°8'N, 
110°58'W,  13.Lx.1966  (Jean  &  Wilton  Ivie,  AMNH); 
20  mi  SW  Sonoyta,  31°38'N,  113°4'W,  13.vi.l952  (W 
J.  Gertsch,  AMNH);  22  mi  E  Hermosillo  on  the 
banks  of  Rio  Sonora,  29°4'N,  110°55'W,  17.viii.l959 
(B.  A.  Branson,  AMNH);  Los  Angeles,  29°27'N, 
110°46'W,  4.X.1966  (V.  Roth,  AMNH);  Puerto  Pefi- 
asco  at  seashore,  31°20'N,  113°33'W,  3.iv.l968  (D.  E. 
Bixler,  MCZ).  Zacatecas:  14  mi  S  Fresnillo,  23°5'N, 
102°45'W,  4.viii.l954  (W  J.  Gertsch,  AMNH);  4  mi 
NE  Concepcion  del  Oro,  24°41'N,  101°23'W, 
4.vii.l984  (J.  B.  Woolley  AD).  USA  Anzona:  Santa 
Catalina  Mtns.,  mile  7.9  of  Catalina  Highway  from 
Tuscon  to  Mt.  Lemmon.,  32°17'N,  110°48'W, 
19.vi.l985  (W  Maddison,  MCZ);  Tucson,  32°15'N, 
110°57'W,  5.vii.l991  (W  H.  Piel  &  G.  S.  Bodner, 
MCZ).  California:  Corn  Springs,  Chuckawalla  Mtns., 
10  mi  SE  Desert  Center,  33°33'N,  115°24'W, 
29.xi.1963  (D.  C.  Lorn,  MCZ);  Manzanita  chaparral, 
San  Gabriel  Canyon,  Coldbrook  Ranger  Station, 
34°11'N,  117°53'W,  19.viii.l964  (L.  Pinter,  MCZ). 
Texas:  Big  Bend  Nat'l  Park,  Old  Ranch  House  on  St. 
Elena  Rd.,  29°10'N,  103°30'W,  25.V.I967  (E.  Sabath, 
MCZ). 

27.  Metepeira  atascadero  new  species 
Figures  214-221,  336;  Map  14 

Holottjpe.  Male  from  San  Miguel  de  Allende,  Guana- 
juato, Mexico,  25.x.  1982,  George  Uetz,  in  MCZ. 
The  specific  name  is  a  noun  in  apposition  after  the 
Rancho  Hotel  Atascadero  in  San  Miguel  de  Allen- 
de. This  name  was  coined  by  George  Uetz  when 
he  and  his  colleagues  stayed  in  Rancho  Hotel  Atas- 
cadero while  studying  the  behavioral  ecology  of  this 
species.  This  name  has  been  in  informal  use  in  the 
literature  (e.g.,  Uetz  and  Hodge,  1990). 

Description.  Female  paratype  from  San 
Miguel  de  Allende,  Guanajuato,  Mexico. 
Reddish  brown  carapace,  white  around 
eyes,  faint  light  marks  extend  posteriorly 
behind  lateral  eyes  (Fig.  220).  Legs  yel- 
lowish, ringed  brown  at  distal  ends  of  ar- 
ticles. Femur  I  with  row  of  four  macro- 
setae  on  anterior  side;  three  on  anterov- 
entral  side.  Largest  branches  of  fleur-de- 
lis  pattern  on  dorsal  folium  form  large 
paired  white  spots  (Fig.  220).  Venter  of  ab- 
domen black  with  wide  white  median  line, 
flanked  by  white  U-shape  mark  (Fig.  221). 
Pair  of  small  white  spots  on  either  side  of 
spiracle.  Sternum  black  with  white  poste- 
rior mark  (Fig.  221).  Ratio  of  eye  diame- 
ters: posterior  medians  and  anterior  me- 
dians 1.0,  anterior  laterals  1.6,  posterior 
laterals  1.3.  Anterior  median  eyes  separat- 


Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


ed  by  1.4  diameters,  posterior  median  eyes 
by  0.9,  anterior  median  eyes  separated 
from  anterior  laterals  by  3.7  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.4  their  diameters.  Total  length  7.5 
mm.  Carapace  3.1  mm  long,  2.5  wide. 
First  femur  4  mm,  patella  and  tibia  4, 
metatarsus  3.9,  tarsus  1.3.  Second  patella 
and  tibia  3.3  mm,  third  1.8,  fourth  2.8. 

Male  holotype.  Reddish  brown  cara- 
pace, lighter  around  eyes,  faint  light  marks 
extend  posteriorly  behind  lateral  eyes  and 
median  eyes  (Fig.  218).  Legs  yellowish 
white,  each  article  gradually  turning  red- 
dish brown  distally.  Femur  I  with  row  of 
four  or  five  macrosetae  on  anterior  side; 
eight  or  nine  on  anteroventral  side.  Larg- 
est branches  of  fleur-de-lis  pattern  on  dor- 
sal folium  form  large  paired  white  spots 
(Fig.  218).  Venter  of  abdomen  black  with 
wide  white  median  line,  flanked  by  faint 
white  U-shape  mark  (Fig.  219).  Pair  of 
small  white  spots  on  either  side  of  spiracle. 
Sternum  black  with  white  posterior  mark 
(Fig.  219).  Ratio  of  eye  diameters:  poste- 
rior medians  and  anterior  medians  1.0,  an- 
terior laterals  1.7,  posterior  laterals  1.4. 
Anterior  median  eyes  separated  by  1.8  di- 
ameters, posterior  median  eyes  by  1.0,  an- 
terior median  eyes  separated  from  anterior 
laterals  by  3.6  diameters  of  anterior  lateral 
eyes,  lateral  eyes  separated  by  0.5  their  di- 
ameters. Total  length  5  mm.  Carapace  2.5 
mm  long,  1.8  wide.  First  femur  4  mm,  pa- 
tella and  tibia  4.2,  metatarsus  4.5,  tarsus 
1.4.  Second  patella  and  tibia  3.3  mm,  third 
1.7,  fourth  2.5. 

Diagnosis.  The  male  embolus  of  M. 
atascadero  has  a  protruding  bump  be- 
tween the  embolus  tip  and  the  basal  em- 
bolic apophysis  (Figs.  214,  217)  which  sep- 
arates it  from  all  other  Metepeira  species. 
As  with  M.  incrasssata  (Fig.  224)  and,  to 
a  lesser  degree,  M.  triangularis  (Fig.  201), 
the  epigynal  depressions  on  either  side  of 
the  scape  of  M.  atascadero  (Fig.  216)  are 
sclerotized  to  create  a  scooped  and  slick 
quality.  The  darker  epigynal  openings  in- 
side the  depressions  are  hidden  anteriorly 
under  the   scapes  hood.   Metepeira  atas- 


cadero's  epigynum  differs  from  that  of  M. 
incrassata  by  its  much  smaller  depressions 
(compare  Fig.  216  with  224). 

Variation.  Average  body  length  of  12  fe- 
males examined  7.9  mm,  range  6  to  9.5 
mm.  Average  body  length  of  12  males  ex- 
amined 5.7  mm,  range  3.5  to  6.9  miTi. 

Natural  History.  Mature  specimens 
have  been  collected  from  the  end  of  Au- 
gust to  the  end  of  October  (Fig.  336).  This 
species  lives  solitarily:  only  20%  of  spiders 
live  in  aggregations  of  two  or  more  indi- 
viduals (Uetz  and  Hodge,  1990).  It  is 
thought  that  the  low  level  of  social  behav- 
ior in  M.  atascadero  occurs  because  the 
species  pursues  a  risk-prone  foraging  strat- 
egy (Uetz,  1988a,b).  Spiders  are  found  be- 
tween 1,500  and  2,500  m  elevation. 

Distribution.  Mexican  highlands  from 
Durango  to  Guerrero  (Map  14). 

Records  Examined.  MEXICO  Coahuila:  Saltillo,  14 
mi  E  in  Larrea  Desert,  25°25'N,  100°55'W, 
28.vii.1944  (AMNH).  Durango:  El  Taseate,  26°12'N, 
105°7'W,  27.vii.1947  (W.  J.  Gertsch,  AMNH);  Yer- 
banis,  80  mi  NW  Durango,  24°45'N,  103°50'W, 
19.viii.l947  (W.  J.  Gertsch,  AMNH).  Guanajuato:  20 
mi  E.  Guanajuato,  21°1'N,  100°57'W,  15.ix.l976  (C. 

E.  Griswold  &  Jackson,  CAS);  Guanajuato,  21°1'N, 
101°15'W  (N.  Banks,  MCZ);  San  Miguel  de  Allende, 
20°55'N,  100°45'W,  25.x.  1982  (George  Uetz,  MCZ), 
14.X.1983  (MCZ),  14.X.1985  (MCZ),  17.x.l98^ 
(MCZ),  26.X.1985  (MCZ).  Guerrero:  Tecalpuico,  25 
km  N  Iguala,  18°29'N,  99°38'W,  l.i.l948  (AMNH). 
Hidalgo:  Ozumbilla,  20°9'N,  101°16'W,  2.X.1957  (R. 
Dreisbach,  MCZ).  Jalisco:  Cyarco  Onda,  30  km  W 
Ojuelos,  21°47'N,  101°53'W  (H.  Wagner,  AMNH). 
Michoacan:  Hills  N  of  Patzcuaro,  19°45'N,  101°36'W, 
24.viii.1959  (A.  F.  Archer,  AMNH);  Hwy  110,  4  mi 
W  Jiquilpan,  19°59'N,  102°47'W,  2.viii.l967  (R.  E. 
Leech,  REL).  Zacatecas:  East  of  Guadalupe, 
22°46'N,  102°31'W,  21.viii.l959  (A.  F.  Archer, 
AMNH);  Guadalupe,  22°45'N,  102°31'W,  16.viii.l947 
(W.  J.  Gertsch,  AMNH). 

28.  Metepeira  incrassata 

F.  O.  P. -Cambridge 

Figures  222-228,  323;  IVIap  9 

Epeira  Salei  KeyserUng,  1864:  93,  fig.,  6,9.  Female 
holotype  from  Oaxaca,  Mexico,  in  BMNH,  exam- 
ined. Keyserhng,  1892:  196,  fig.  145,  9.  Roewer, 
1942:  851.  NEW  SYNONYMY. 

Metepeira  incrassata  F.  O.  R-Cambridge,  1904:  460, 
fig.  11,  ?.  Female  holotype  from  Jalapa,  Mexico, 
in  BMNH.  Roewer,  1942:  868.  Bonnet,  1957:  2821. 


Metepeira  •  Piel 


69 


220"^"      221 

atascadero 

(27) 


224 


^im^  227   '        228 

incrassata 
(28) 


234 


Figures  214-221.  Metepeira  atascadero  new  species  (sp.  27;  20°55'N,  100°45'W).  214,  male  palpus,  mesal.  215,  epigynum, 
posterior.  216,  epigynum,  ventral.  217,  male  embolic  division,  ventral.  218,  male,  dorsal.  219,  male,  ventral.  220,  female,  dorsal. 

221,  female,  ventral. 

Figures  222-228.  Metepeira  incrassata  F.  O.  P.-Cambridge  (sp.  28  [222,225-228]  19°4'N,  97°2'W;  [223,224]  18°54'N,  97°0'W). 

222,  male  palpus,  mesal.  223,  epigynum,  posterior.  224,  epigynum,  ventral.  225,  male,  dorsal.  226,  male,  ventral.  227,  female, 
dorsal.  228,  female,  ventral. 

Figures  229-235.  Metepeira  ventura  Chamberlin  and  Ivie  (sp.  29;  28°5'N,  114°8'W).  229,  male  palpus,  mesal.  230,  epigynum, 
posterior.  231,  epigynum,  ventral.  232,  male,  dorsal.  233,  male,  ventral.  234,  female,  dorsal.  235,  female,  ventral. 
Scale  bars:  dorsum  and  venter  figures  1.0  mm. 


70 


Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No. 


Aranea  sallei: — F.O.P.-Cambridge,  1904:  519.  Roew- 
er,  1942:  851. 

Araneus  incrassata: — Petrunkevitch,  1911:  289. 

Araneus  sallei: — Petrunkevitch,  1911:  314. 

Metepeira  salei: — Levi,    1991:    180.    Platnick,    1993: 
449. 

Note.  Although  M.  incrassata  is  not  the  oldest 
name,  it  has  been  cited  more  tlian  20  times  in  the 
general  literature  (e.g.,  Caraco  et  al.  1995;  Hieber 
and  Uetz,  1990;  Hodge  and  Uetz,  1992,  1995, 
1996;  Jakob  et  al.,  1996;  Rayor,  1996;  Rayor  and 
Uetz,  1990,  1993;  Uetz,  1988a,b,  1989,  1991,  1992, 
1996;  Uetz  and  Hieber,  1994,  1997;  Uetz  and  Hod- 
ge, 1990;  Uetz  et  al.,  1994).  In  contrast,  the  senior 
synonym  has  not  been  cited  outside  of  taxonomic 
catalogues.  In  compliance  with  Article  79  of  the 
ICZN  (1985),  I  choose  to  assert  the  priority  of  tlie 
more  popular  junior  name. 

Description.  Female  from  Rancho  Chu- 
la-Vista,  Veracruz,  Mexico.  Browii  cara- 
pace, darker  around  margins,  lighter 
around  eyes  and  behind  lateral  eyes  (Fig. 
227).  Legs  brown,  yellow  at  base  of  articles 
and  on  articles  distal  to  the  femur  for  legs 
I  and  II.  Femur  I  with  row  of  three  to  four 
macrosetae  on  anterior  side;  one  on  anter- 
oventral  side.  Dorsal  folium  darker  than 
most  Metepeira  species.  White  fleur-de-lis 
pattern  with  thin  branches  (Fig.  227).  Ven- 
ter dark  brown  to  black  with  white  median 
mark  that  is  shorter  than  it  is  in  most  other 
species  (Fig.  228).  Sternum  dark  brown  to 
black  with  one  or  two  small  white  spots, 
usually  in  the  center  or  anteriorly  (Fig. 
228).  Ratio  of  eye  diameters:  posterior  me- 
dians and  anterior  medians  0.9,  anterior 
laterals  1.3,  posterior  laterals  1.1.  Anterior 
median  eyes  separated  by  1.5  diameters, 
posterior  median  eyes  by  1.1,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  4.1  diameters  of  anterior  lateral  eyes, 
lateral  eyes  separated  by  0.5  their  diame- 
ters. Total  length  7.8  mm.  Carapace  3.6 
mm  long,  2.9  wide.  First  femur  3.9  mm, 
patella  and  tibia  4.1,  metatarsus  3.5,  tarsus 
1.4.  Second  patella  and  tibia  3.7  mm,  third 
2.3,  fourth  3.3. 

Male  from  Rancho  Chula-Vista,  Vera- 
cruz, Mexico.  Carapace,  dorsum,  venter, 
sternum  a  darker  version  of  female  (Figs. 
225,  226).  Base  of  femora  yellow,  remain- 
der dark  brown;   other  articles  gradually 


turning  lighter  distally  Femur  I  with  row 
of  four  to  six  macrosetae  on  anterior  side; 
six  to  nine  on  anteroventral  side.  Ratio  of 
eye  diameters:  posterior  medians  and  an- 
terior medians  1.0,  anterior  laterals  1.4, 
posterior  laterals  1.2.  Anterior  median 
eyes  separated  by  1.4  diameters,  posterior 
median  eyes  by  1.0,  anterior  median  eyes 
separated  from  anterior  laterals  by  3.7  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.4  their  diameters.  To- 
tal length  7  mm.  Carapace  3.3  mm  long, 
2.7  wide.  First  femur  4.9  mm,  patella  and 
tibia  5.3,  metatarsus  4.8,  tarsus  1.7.  Sec- 
ond patella  and  tibia  4.3  mm,  third  2.2, 
fourth  3.4. 

Diagnosis.  Overall  pigmentation  of  M. 
incrassata  is  darker  than  all  others  in  its 
species  group.  The  thin  branches  on  the 
dorsal  folium  (Figs.  225,  227),  the  almost 
entirely  dark  sternum,  and  the  shortened 
median  line  on  the  venter  (Figs.  226,  228) 
are  distinctive.  As  with  M.  ohnec  (Fig. 
178),  M.  comanche  (Fig.  185),  and  M.  tri- 
angularis (Fig.  199),  the  embolus  of  M.  in- 
crassata (Fig.  222)  curves  off  sharply  from 
a  distal  embolic  apophysis  that  does  not 
project  forward.  Compared  to  M.  olniec 
(Fig.  178)  and  M.  comanche  (Fig.  185),  M. 
incrassata's  embolus  tip  beyond  the  distal 
apophysis  is  relatively  longer  and  not  as 
curved  (Fig.  222).  Male  M.  incrassata  are 
easily  separated  from  M.  triangularis  by 
the  shape  of  the  median  apophysis  (com- 
pare Fig.  222  with  Fig.  199).  As  with  M. 
atascadero  (Fig.  216)  and,  to  a  lesser  de- 
gree, M.  triangularis  (Fig.  201),  the  epi- 
gynal  depressions  on  either  side  of  the 
scape  of  M.  incrassata  (Fig.  224)  are  scler- 
otized  to  create  a  scooped  and  slick  qual- 
ity. The  darker  epigynal  openings  inside 
the  depressions  are  hidden  anteriorly  un- 
der the  scape's  hood.  Metepeira  incrassa- 
ta's  epigynum  differs  from  those  of  M. 
atascadero  and  M.  triangularis  by  its  much 
larger  and  disk-shaped  depressions  (com- 
pare Fig.  224  with  Figs.  201,  216). 

Variation.  Average  body  length  of  31  fe- 
males examined  7.6  mm,  range  6.4  to  9.1 


Metepeira  •  Piel 


71 


mm.  Average  body  length  of  22  males  ex- 
amined 6.2  mm,  range  4.3  to  8  mm. 

Natural  History.  Mature  specimens 
have  been  collected  in  February  through 
October  but  are  most  frequently  found  in 
July  (Fig.  323).  This  species  lives  socially: 
half  of  all  spiders  live  in  aggregations  of 
1,000  or  more  individuals  (Uetz  and  Hod- 
ge, 1990).  Colonies  are  frequently  found 
spanning  telephone  lines,  houses,  fences, 
and  other  man-made  structures.  It  is 
thought  that  this  high  level  of  social  be- 
havior occurs  because  the  species  pursues 
a  risk-averse  foraging  strategy  in  which  the 
variance  in  survival  attenuates  with  in- 
creasing colony  size  (Uetz,  1988a,b).  Ele- 
vations center  around  800  m  and  range 
from  about  500  to  1,500  m. 

Distribution.  Tropical  Mexico  from  San 
Luis  Potosi  to  southern  Veracruz  and  Oa- 
xaca  (Map  9). 

Records  Examined.  MEXICO  Hidalgo:  20  mi  S  of 
Jacala,  20°50'N,  99°16'W,  18.iv.l946  (L.  I.  Davis  & 
M.  Johnston,  AMNH);  Chapulhuacan,  21°10'N, 
98°54'W,  20.V.1952  (M.  Cazier,  W.  Gertsch,  &  R. 
Schrammel,  AMNH),  27.vii.1966  (Jean  &  Wilton 
Ivie,  AMNH),  16.vii.l969  (S.  &  J.  Peck,  MCZ).  Mex- 
ico: 1  mi  S  Palomas,  19°50'N,  99°5'W,  28.vii.1950 
(AMNH).  Oaxaca:  Oaxaca,  17°3'N,  96°43'W  (Nathan 
Banks,  MCZ).  Puebla:  Huauchinango,  20°11'N, 
98°3'W,  7..X.1947  (H.  Wagner,  AMNH);  north  of  Xic- 
otepec  de  Juarez,  20°18'N,  97°57'W  19.ivl967  (W 
B.  Peck,  MCZ).  San  Luis  Potosi:  10km  W  Xilitla  on 
rl20,  21°22'N,  99°4'W  10.viii.l991  (W  H.  Piel  &  G. 
S.  Bodner,  MCZ).  Veracruz:  1  mi  SW  Tlapacoyan, 
19°57'N,  97°14'W  16.vii.l973  (A.  Newton,  MCZ);  2 
mi  N  Fortin  de  las  Flores,  18°56'N,  97°1'W, 
5.viii.l966  (Jean  &  Wilton  Ivie,  AMNH);  5  mi  E  Ori- 
zaba, 18°51'N,  97°4'W  25.vii.1956  (W  Gertsch  &  V. 
Roth,  AMNH);  Coatepec,  19°27'N,  96°58'W, 
28.vii.1955  (C.  &  P  Vaurie,  AMNH),  19.vii.l991  (W. 
H.  Piel  &  G.  S.  Bodner,  MCZ);  Fortin  de  las  Flores, 
18°54'N,  97°0'W,  7.vii.l947  (G.  &  M.  Goodnight, 
AMNH),  25.iv.1963  (W.  J.  Gertsch  &  W  Ivie, 
AMNH),  10.vii.l976  (A.  Newton,  MCZ),  21.X.1982 
(George  Uetz,  MCZ),  17.vii.l991  (W.  H.  Piel  &  G.  S. 
Bodner,  MCZ);  Jalapa,  19°32'N,  96°55'W,  14.ii.l948 
(H.  Wagner,  AMNH),  15.iii.l948  (H.  Wagner, 
AMNH),  15.X.1962  (N.  L.  H.  Krauss,  AMNH);  Los 
Naranjos,  18°21'N,  96°10'W  4.iii.l948  (H.  Wagner, 
AMNH);  near  Monte  Blanco,  18°58'N,  97°1'W 
3.viii.l973  (A.  Newton,  MCZ);  Orizaba,  18°51'N, 
97°4'W,  6.vii.l963  (D.  BLxler,  MCZ);  Rancho  Chula- 
Vista,  N  of  Cordoba,  19°4'N,  97°2'W  18.vii.l991  (W 
H.  Piel  &  G.  S.  Bodner,  MCZ). 


Metepeira  ventura  Group 

Spiders  in  the  M.  ventura  group  {Me- 
tepeira ventura,  Metepeira  revillagigedo , 
Metepeira  celestun,  Metepeira  uncata,  Me- 
tepeira crassipes,  Metepeira  chilapae)  are 
closely  related  to  those  in  the  M.  minima 
group  (M.  petatlan,  M.  minima,  M.  paci- 
fica,  M.  jamaicensis) .  Females  in  the  M. 
ventura  group  have  epigynal  openings  that 
are  wider  than  long  and  shaped  as  trian- 
gles (Fig.  231),  ovals  (Fig.  252),  or  squares 
(Figs.  259,  266),  with  their  posterior  edges 
open.  The  posterior  edges  are  observed  to 
be  open  because  the  posterior  lobes  at 
their  distal  edge  are  wider  than  the  distal 
end  of  the  scape.  In  many  other  species, 
the  gap  between  the  lobes  is  narrower 
than  the  scape  and  therefore  hidden  from 
a  ventral  view,  giving  the  impression  that 
the  openings  form  closed  shapes  (e.g..  Fig. 
123).  In  addition,  with  the  exception  of  M. 
celestun  (Fig.  245),  the  scape  is  relatively 
thin  throughout  its  entire  length  (Figs. 
231,  238,  252,  259,  266).  In  males,  the  em- 
bolus can  be  slini  and  elongated  with  a 
gentle  curve  (e.g..  Figs.  236,  243),  or  as 
with  some  species  in  the  M.  minima  group, 
it  can  be  tapering  to  a  shaip  bend  right  at 
the  tip  (e.g..  Figs.  229,  264).  In  general, 
the  larger  flagellum  is  a  simple  tapering 
extension  off  the  base  (Figs.  229,  243,  250, 
257,  264,  and  to  a  lesser  extent,  Fig.  236), 
as  opposed  to  being  a  distinctly  separate 
structure  that  abruptly  cuives  off  the  base 
(e.g..  Fig.  129).  Although  the  flagellae  can 
be  thin,  as  in  the  M.  minima  species 
group,  they  are  not  set  off  on  a  distinctly 
narrower  stalk  (e.g.,  compare  Fig.  264  with 
Fig.  278). 

29.  Metepeira  ventura 
Chamberlin  and  Ivie 
Figures  229-235,  320;  Map  13 

Metepeira  ensenada  Chamberlin  and  Ivie,  1942:  65, 
figs.  166-168,  6.  Male  holotype  from  beach  near 
Ensenada,  Mexico,  in  the  AMNH,  examined.  Syn- 
onymized  by  Levi  (1977). 

Metepeira  ventura  Chamberlin  and  Ivie,  1942:  67, 
figs.  175-179,  ?.  Female  holotype,  1  male  and  3 
female  paratypes  from  between  Oxnard  and  Santa 
Monica,  California,  USA,  in  the  AMNH,  examined. 


72         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


Levi,  1977:  204,  figs.  53-60.  Brignoli,  1983:  275. 

Note.  As  first  reviser,  Levi  (1977)  preferred  to 
use  the  name  M.  ventura  because  a  larger  number 
of  specimens  were  available  from  the  type  locality. 

Description.  Female  from  10  mi  north 
of  Colonia  Guerrero,  Baja  California  Nor- 
te, Mexico.  Carapace  brown  with  white 
eye  region,  lateral  posterior  extensions, 
and  white  median  line  (Fig.  234).  Legs  yel- 
lowish tan  with  darker  rings  on  fourth  pair 
of  legs.  Femur  I  with  row  of  four  macro- 
setae  on  anterior  side;  two  on  anteroven- 
tral  side.  Dorsum  of  abdomen  with  usual 
Metepeira  folium,  lighter  in  anterior  third 
(Fig.  234).  Venter  of  abdomen  with  wide 
white  median  line  posteriorly  set  within 
hint  of  surrounding  U-shaped  marking. 
Pair  of  white  spots  on  either  side  of  spi- 
racle (Fig.  235).  Sternum  has  wide  inedian 
white  line  widening  anteriorly,  sometimes 
fragmented  (Fig.  235).  Ratio  of  eye  di- 
ameters: posterior  medians  and  anterior 
medians  1.0,  anterior  laterals  1.6,  posterior 
laterals  1.4.  Anterior  median  eyes  separat- 
ed by  1.4  diameters,  posterior  median  eyes 
by  0.7,  anterior  median  eyes  separated 
from  anterior  laterals  by  2.9  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.3  their  diameters.  Total  length  5.5 
mm.  Carapace  2.8  mm  long,  2.2  wide. 
First  femur  3.3  mm,  patella  and  tibia  3.3, 
metatarsus  3,  tarsus  1.2.  Second  patella 
and  tibia  2.9  mm,  third  1.7,  fourth  2.5. 

Male  from  10  mi  north  of  Colonia 
Guerrero,  Baja  California  Norte,  Mexico. 
Carapace,  abdomen,  venter  as  in  female 
(Figs.  232,  233).  Ringed  legs  darker  than 
in  female.  Femur  I  with  row  of  four  ma- 
crosetae  on  anterior  side;  six  to  seven  on 
anteroventral  side.  Median  white  line  on 
sternum  often  broken  (Fig.  233).  Ratio  of 
eye  diameters:  posterior  medians  and  an- 
terior medians  0.9,  anterior  laterals  1.2, 
posterior  laterals  1.1.  Anterior  median 
eyes  separated  by  1.6  diaineters,  posterior 
median  eyes  by  0.8,  anterior  median  eyes 
separated  from  anterior  laterals  by  1.9  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.3  their  diameters.  To- 
tal length  4.3  mm.  Carapace  2.2  mm  long, 


1.7  wide.  First  femur  3.6  mm,  patella  and 
tibia  3.6,  metatarsus  3.7,  tarsus  1.3.  Sec- 
ond patella  and  tibia  2.8  mm,  third  1.5, 
fourth  2.3. 

Diagnosis.  Unlike  other  members  of  the 
M.  minima  species  group,  the  larger  fla- 
gellum  on  the  median  apophysis  of  M. 
ventura  is  a  tapering  extension  off  the  base 
(Fig.  229),  as  opposed  to  arising  from  a 
distinctly  separate  stalk  (Fig.  286).  Like  M. 
uncata,  the  epigynal  openings  of  M.  ven- 
tura (Fig.  231)  are  somewhat  triangular  in 
shape,  in  contrast  to  oval  (Fig.  288)  or  slit- 
shaped  (Fig.  280)  as  in  other  members  of 
the  M.  minima  species  group.  The  rela- 
tively shorter  scape  and  the  dark  circles 
inside  the  epigynal  openings  of  M.  ventura 
(Fig.  231)  contrast  with  M.  uncata  s  rela- 
tively longer  scape  and  the  dark  sinuous 
shapes  inside  the  epigynal  openings  (Fig. 
252). 

Variation.  Average  body  length  of  eight 
females  examined  6.7  mm,  range  5.6  to  9.7 
mm.  Average  body  length  of  seven  males 
examined  5.1  mm,  range  3.9  to  7  mm. 

Natural  History.  Mature  speciinens  are 
mostly  collected  April  through  September 
(Fig.  320).  Altitudes  range  from  near  sea 
level  to  1,000  m. 

Distribution.  California  to  northwestern 
Mexico  (Levi,  1977,  map  1;  Map  13). 

Records  Examined.  MEXICO  Baja  Calif.  Not~te:  10 
m  E  El  Rosario,  30°11'N,  115°46'W,  8.vii.l973  (S.  C. 
Wilhams  &  K.  B.  Blair,  CAS);  10  mi  N  Col.  Guerrero, 
28°5'N,  114°8'W,  l.b(.1957  (V.  Roth,  AMNH);  2  mi 
SE  Erendira,  3ri9'N,  116°19'W,  12.V.1973  (S.  C. 
Wilhams  &  K.  B.  Blair,  CAS);  Isla  Cedros,  Gran  Cafi- 
on,  28°12'N,  115°15'W,  10.iii.l945  (B.  F.  Osorio  Taf- 
all,  AMNH);  Isla  de  Cedros,  Cerro  de  Cedros, 
28°12'N,  115°15'W,  l.vii.l983  (V.  F.  Lee,  CAS);  Isla 
de  Cedros,  Punta  Norte  [?],  28°22'N,  115°14'W, 
3.vii.l983  (V.  F.  Lee,  CAS);  Isla  de  Cedros,  trail  to 
Cerro  de  Cedros,  at  spring,  28°12'N,  115°15'W, 
27.Lx.1984  (D.  B.  Weissman,  V.  F  Lee,  CAS);  Islas 
San  Benito,  Benitos  del  Oeste,  28°18'N,  115°35'W, 
4.vii.l983  (D.  C.  Lightfoot  &  V.  F  Lee,  CAS);  Islas 
San  Benito,  Middle  Island,  28°19'N,  115°34'W, 
9.iv.l981  (Stanley  C.  Williams,  CAS);  Islas  San  Be- 
nito, South  Island,  28°18'N,  115°35'W,  9.iv.l981 
(Stanley  C.  Williams,  CAS);  near  Consuelo,  6  mi  NW 
El  Rosario,  30°11'N,  115°46'W  18.iv.l965  (D.  Q. 
Cavagnaro,  C.  E.  &  E.  S.  Ross,  V.  L.  Vesterby,  CAS); 
Rancho  Las  Parritas,  10  mi  S  San  Quintin,  30°20'N, 


Metepeira  •  Piel 


73 


115°57'W,  27.vi.1977  (C.  E.  Griswold,  CAS);  Santo 
Tomas,  31°33'N,  116°24'W,  8.vii.l953  (W.  J.  &  J.  W. 
Gertsch,  AMNH).  Baja  Calif.  Sun  Bahia  de  los 
Muertos,  23°58'N,  109°50'W,  20.xii.l958  (H.  B. 
Leech,  CAS);  Desierto  del  Vizcaino,  Laguna  Ojo  de 
Liebre,  sobre  frutilla,  27°43'N,  114°15'W,  ll.xi.l981 
(A.  Cota  &  M.  Jimenez,  MLJ);  E  edge  of  Sierra  Pla- 
ceres,  27°35'N,  114°30'W,  25.iii.1984  (W.  J.  Pulawsld, 
CAS).  Sonora:  6  mi  E  Navojoa,  27°6'N,  109°23'W, 
23.viii.1965  (W.  J.  Gertsch  &  R.  Hastings,  AMNH). 
USA  California:  1  mi  NW  Winchester  (Double 
Butte),  33°43'N,  117°6'W,  7.xii.l976  (W.  Icenogle, 
MCZ);  Lompoc,  by  US  63,  34°38'N,  120°27'W, 
17.viii.l966  (L.  &  P.  Pinter,  MCZ);  Lucia,  Monterey 
Co.,  36°1'N,  121°33'W,  15.ix.l964  (L.  Pinter,  MCZ); 
Manzanita  chaparral,  San  Gabriel  Canyon,  Coldbrook 
Ranger  Station,  34°11'N,  117°53'W,  29.V.1965  (L. 
Pinter,  MCZ);  Santa  Catalina  Island  Area  near  Hay- 
press  Res,  33°23'N,  118°25'W,  6.ii.l993  (Martin  C. 
Ramirez  &  Laura  B.  Fandino,  MCZ);  Santa  Catalina 
Island,  Mt.  Torquemada,  33°26'N,  118°33'W, 
15.viii.l965  (L.  Pinter,  MCZ);  Winchester,  33°42'N, 
117°5'W,  ll.v.1970  (W.  Icenogle,  MCZ);  Winchester, 
Double  Butte,  33°42'N,  117°5'W,  19.V.1974  (W.  Icen- 
ogle, MCZ).  Nevada:  Quin  River  Crossing,  41°35'N, 
118°27'W,  21.vi.l975  (G.  F.  Knowlton,  MCZ). 

30.  Metepeira  revillagigedo  new  species 
Figures  236-242,  301;  IVIap  12 

Holotype.  Female  from  south  of  Isla  Socorro,  Archi- 
pielago  de  Revillagigedo,  State  of  Colima,  Me.xico, 
15.xii.l988,  M.  Jimenez,  in  MCZ.  The  specific 
name  is  a  noun  in  apposition  after  the  locality. 

Description.  Female  holotype.  Carapace 
dark  brown  with  light  region  in  median 
eye  quadrangle  and  surrounding  lateral 
eyes.  Pair  of  darker,  elliptic,  walnut  leaf 
shapes  on  lighter  patch  near  center  of  car- 
apace (see  Fig.  241).  Darker  rings  on  distal 
ends  of  femora,  patellae,  tibiae,  and  prox- 
imal dorsal  portion  of  tibia  I,  II.  Femur  I 
with  three  to  four  macrosetae  on  anterior 
side;  none  on  the  anteroventral  side.  Dor- 
sum of  abdomen  with  white  oak  leaf  foli- 
um, margined  with  dark  markings,  partic- 
ularly on  trailing  edges  of  lobes  (Fig.  241). 
Coloration  has  a  slightly  golden  hue.  Dark 
lateral  band  wraps  around  sides  of  abdo- 
men and  stretches  up  over  anterior  dorsal 
portion.  Venter  with  two  white  spots  on 
either  side  of  spiracle.  Anterior  to  spiracle, 
slight  indication  of  a  V-shaped  mark  with 
longitudinal  extensions  reaching  halfway 
up  abdomen.  Wide,  white  medial  longitu- 


dinal line  to  epigynal  groove  (Fig.  242). 
Sternum  with  wide,  white  longitudinal 
mark  widening  anteriorly  (Fig.  242).  Ratio 
of  eye  diameters:  posterior  medians  and 
anterior  medians  0.8,  anterior  laterals  1.1, 
posterior  laterals  1.  Anterior  median  eyes 
separated  by  1.3  diameters,  posterior  me- 
dian eyes  by  1.0,  anterior  median  eyes  sep- 
arated from  anterior  laterals  by  2.5  diam- 
eters of  anterior  lateral  eyes,  lateral  eyes 
separated  by  0.3  their  diameters.  Total 
length  7.5  mm.  Carapace  3.2  mm  long,  2.5 
wide.  First  femur  3.5  mm,  patella  and  tibia 
3.8,  metatarsus  3.2,  tarsus  1.2.  Second  pa- 
tella and  tibia  3.2  mm,  third  1.8,  fourth 
2.8. 

Male  paratype  froin  Isla  Socorro,  Coli- 
ma. Carapace  dirty  yellowish  brown  with 
lighter  posteriorly  pointing  acute  triangle 
in  center  (Fig.  239).  Legs  same  color  as 
carapace,  except  lighter  on  proximal  half 
of  femora.  Femur  I  with  row  of  four  ma- 
crosetae on  anterior  side,  row  of  two  ma- 
crosetae on  anteroventral  side.  Male  ab- 
domen lighter  and  with  less  contrast  than 
female  (Fig.  239).  Venter  with  a  much  re- 
duced and  shorter  longitudinal  line  than 
female.  Sternum  as  in  female,  except  lon- 
gitudinal line  more  often  broken  (Fig. 
240).  Ratio  of  eye  diameters:  posterior  me- 
dians and  anterior  medians  1.0,  anterior 
laterals  1.1,  posterior  laterals  1.1.  Anterior 
median  eyes  separated  by  1.5  diameters, 
posterior  median  eyes  by  0.8,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  1.4  diameters  of  anterior  lateral  eyes, 
lateral  eyes  separated  by  0.3  their  diame- 
ters. Total  length  3.2  mm.  Carapace  1.7 
mm  long,  1.1  wide.  First  femur  2.5  mm, 
patella  and  tibia  2.5,  metatarsus  2.3,  tarsus 
0.9.  Second  patella  and  tibia  2  mm,  third 
1.0,  fourth  1.5. 

Diagnosis.  The  epigynum  of  M.  revilla- 
gigedo looks  veiy  different  from  those  of 
other  species  in  the  M.  ventura  group.  In- 
stead of  having  small,  sharply  delineated 
depressions  on  either  side  of  the  scape 
(e.g..  Fig.  259),  M.  revillagigedo  has  larger 
but  more  gradual  depressions  (Fig.  238). 
Despite  the  unique  appearance  of  the  epi- 


74         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


gynum,  the  affinity  between  M.  revillagi- 
gedo  and  the  M.  ventura  species  group  can 
be  seen  in  the  black  comma-shaped  marks 
inside  the  epigynal  depressions  (Fig.  238) 
similar  to  those  in,  for  example,  M.  Ven- 
tura (Fig.  231).  The  male  emboli  of  M. 
revillagigedo  (Fig.  236)  and  M.  celestun 
(Fig.  243)  are  veiy  similar  in  shape  and 
relatively  longer  than  in  other  species  in 
the  M.  Ventura  group.  Males  of  M.  revil- 
lagigedo differ  from  males  of  M.  celestun 
by  the  thickness  and  shape  of  the  median 
apophysis  (compare  Fig.  236  with  Fig. 
243). 

Natural  History.  Specimens  have  only 
twice  been  collected:  once  in  December 
and  once  in  April  (Fig.  301).  This  species 
is  known  to  live  in  fig  trees. 

Distribution.  This  species  is  found  on 
Isla  Socorro  in  the  Pacific  Ocean  (Map 
12),  the  island  being  one  of  several  that 
form  the  Archipielago  de  Revillagigedo.  It 
is  well  isolated,  situated  about  465  km 
south  of  Baja  California  Sur  and  588  km 
west  of  the  Jalisco  coastline. 

Records  Examined.  MEXICO  Colinia:  Archipiela- 
go de  Revillagigedo:  Isla  Socorro,  18°45'N,  110°57'W, 
26.iv.1932  (Templeton,  Crocker  Exped.,  CAS);  Ar- 
chipielago de  Revillagigedo:  Sur  de  la  Isla  Socorro, 
18°44'N,  110°57'W,  15.xii.l988  (M.  Jimenez,  MCZ). 

31 .  Metepeira  celestun  new  species 
Figures  243-249,  326;  IVlap  13 

Holotype.  Male  from  Celestun,  Yucatan,  Mexico, 
24.vii.1991,  W.  H.  Piel  &  G.  S.  Bodner,  in  MCZ. 
The  specific  name  is  a  noun  in  apposition  after  the 
locality. 

Note.  Since  males  of  this  species  were  never 
found  witli  females,  one  cannot  be  sure  absolutely 
that  they  are  conspecific.  Several  facts  argue  for 
conspecificity:  all  specimens  were  collected  in  sim- 
ilar habitats  and  during  the  same  season;  dorsal  fo- 
lium of  males  and  females  share  similar  patterns 
and  both  have  hint  of  gold  coloration;  based  on 
parallel  correspondence  widi  other  species,  die 
male  palp  is  arguably  compatible  with  the  female  s 
epigynum. 

Description.  Female  paratype  from 
Edzna,  Campeche,  Mexico.  Carapace  yel- 
lowish tan  with  lighter  area  around  eyes 
(Fig.  248).  Legs  yellowish  tan  with  darker 
rings  on  fourth  pair  of  legs.  Femur  I  with 


row  of  four  macrosetae  on  anterior  side; 
sometimes  four  on  anteroventral  side. 
Dorsal  folium  as  in  other  Metepeira,  ex- 
cept speckled  an  unusual  reddish  gold  col- 
or (Fig.  248).  Venter  wide  white  median 
line  with  pair  of  large  white  spots  on  either 
side  of  spiracle  (Fig.  249).  Sternum  has 
wide  median  white  line  widening  anteri- 
orly with  constriction  in  center  (Fig.  249). 
Ratio  of  eye  diameters:  posterior  medians 
and  anterior  medians  1.0,  anterior  laterals 

1.3,  posterior  laterals  1.2.  Anterior  median 
eyes  separated  by  1.5  diameters,  posterior 
median  eyes  by  0.9,  anterior  median  eyes 
separated  froin  anterior  laterals  by  2.4  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.2  their  diameters.  To- 
tal length  5.2  mm.  Carapace  2.5  mm  long, 
1.9  wide.  First  femur  2.6  mm,  patella  and 
tibia  2.8,  metatarsus  2.2,  tarsus  0.9.  Sec- 
ond patella  and  tibia  2.4  mm,  third  1.4, 
fourth  2. 

Male  holotype.  Carapace  dark  with  light 
region  around  eyes  and  a  light  triangular 
mark  anterior  to  thoracic  furrow  (Fig. 
246).  Coxae  and  proximal  third  of  femora 
white,  remaining  two-thirds  black;  other 
articles  similarly  ringed.  Femur  I  with  row 
of  four  macrosetae  on  anterior  side;  three 
on  anteroventral  side.  Dorsal  folium  with 
slight  gold  iridescence.  Fleur-de-lis  pat- 
tern leafy  and  pinnate  (Fig.  246).  Venter, 
sternum  as  in  female,  except  median  white 
line  on  sternum  often  broken  (Fig.  247). 
Ratio  of  eye  diameters:  posterior  medians 
and  anterior  medians  1.1,  anterior  laterals 

1.4,  posterior  laterals  1.3.  Anterior  median 
eyes  separated  by  1.6  diameters,  posterior 
median  eyes  by  0.9,  anterior  median  eyes 
separated  from  anterior  laterals  by  1.4  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.2  their  diameters.  To- 
tal length  2.8  mm.  Carapace  1.4  mm  long, 
1.1  wide.  First  femur  1.8  mm,  patella  and 
tibia  1.8,  metatarsus  1.4,  tarsus  0.8.  Sec- 
ond patella  and  tibia  1.5  mm,  third  0.8, 
fourth  1.2. 

Diagnosis.  The  overall  coloration  of  M. 
celestun  in  alcohol  has  an  unusually  golden 
quality  that  is  rare  among  preserved  Me- 


Metepeira  •  Piel         75 


Figures  236-242.  Metepeira  revillagigedo  new  species  (sp.  30;  18°44'N,  110°57'W).  236,  male  palpus,  mesal.  237,  epigynum, 

posterior.  238,  epigynum,  ventral.  239,  male,  dorsal.  240,  male,  ventral.  241,  female,  dorsal.  242,  female,  ventral. 

Figures  243-249.  Metepeira  celestun  new  species  (sp.  31    [243,246,247]  20°56'N,  90°21'W;  [244,245,248,249]  19°35'N, 

90°15'W).  243,  male  palpus,  mesal.  244,  epigynum,  posterior.  245,  epigynum,  ventral.  246,  male,  dorsal.  247,  male,  ventral. 

248,  female,  dorsal.  249,  female,  ventral. 

Figures  250-256.  Metepeira  uncata  F.  O.  P.-Cambridge  (sp.  32  [250,251,253-256]  14°40'N,  92°9'W;  [252]  14°49'N,  91°31'W), 

250,  male  palpus,  mesal.  251,  epigynum,  posterior.  252,  epigynum,  ventral.  253,  male,  dorsal.  254,  male,  ventral.  255,  female, 

dorsal.  256,  female,  ventral. 

Scale  bars:  dorsum  and  venter  figures  1.0  mm. 


76         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


tepeira.  Unlike  all  other  members  of  the 
M.  Ventura  species  group,  the  scape  of  M. 
celestun  is  very  wide  at  its  base,  thereby 
partly  concealing  the  epigynal  openings 
(Fig.  245).  However,  if  the  scape  is  dis- 
carded, the  exposed  openings  have  black 
comma-shaped  marks  similar  to  those  of 
M.  Ventura  (Fig.  231),  thereby  confirming 
the  species'  affinities  with  other  members 
in  the  M.  ventura  species  group.  Unlike 
other  members,  the  posterior  lobes  nearly 
touch  each  other  near  their  base  (Fig. 
244).  The  emboli  of  M.  celestun  and  M. 
revillagigedo  are  very  similar  in  shape  and 
relatively  longer  than  in  other  species  in 
the  M.  ventura  group  (Fig.  243).  Metepei- 
ra  celestun  differs  from  M.  revillagigedo  by 
the  thickness  and  shape  of  the  median 
apophysis  (compare  Fig.  243  with  Fig. 
236). 

Variation.  Average  body  length  of  four 
feinales  examined  5.6  mm,  range  5.2  to  6.4 
mm.  Average  body  length  of  three  males 
examined  2.7  mm,  range  2.2  to  2.9  mm. 
Female  scape  sometimes  wider  than  it  ap- 
pears in  Fig.  245,  resembling  Metepeira 
arizonica.  Legs  and  carapace  color  vary 
from  yellowish  tan  to  orange-red. 

Natural  History.  Mature  specimens 
have  been  collected  in  July  (Fig.  326)  in 
forested  clearings,  roadside  bushes,  palms, 
and  swampy  areas  near  the  beach. 

Distribution.  Yucatan  peninsula  (Map 
13). 

Records  Examined.  MEXICO  Campeche:  Edzna, 
19°35'N,  90°15'W,  22.vii.1991  (W.  H.  Piel  &  G.  S. 
Bodner,  MCZ).  Quintana  Roo:  Kohunlich  ruins,  9  km 
S  Franciso  Villa,  18°26'N,  88°48'W,  15.vii.l983  (R.  S. 
Anderson,  MCZ).  Yucatan:  3km  S  San  Felipe, 
21°32'N,  88°14'W,  25.vii.1991  (W.  H.  Piel  &  G.  S. 
Bodner,  MCZ);  4  km  N  Xocenpich,  12  km  N  Piste, 
on  road  to  Dzitas,  20°47'N,  88°34'W,  20.vii.l983  (W. 
Maddison,  MCZ);  Balankanche  Cave,  2  km  E  Chi- 
chen  Itza,  20°40'N,  88°33'W,  19.vii.l983  (W.  Mad- 
dison, MCZ);  beach  north  of  Celestun,  20°56'N, 
90°21'W,  24.vii.1991  (W.  H.  Piel  &  G.  S.  Bodner, 
MCZ);  Chichen  Itza  ruins  on  HWY  180,  seasonal  for- 
est, 20°40'N,  88°34'W,  19.vii.l983  (W.  Maddison  & 
R.  S.  Anderson,  MCZ);  Uxmal,  20°22'N,  89°46'W, 
23.vii.1991  (W.  H.  Piel  &  G.  S.  Bodner,  MCZ). 


32.  Metepeira  uncata 
F.  O.  P. -Cambridge 
Figures  250-256,  329;  Map  12 

Metepeira  uncata  F.  O.  P.-Cambridge,  1903:  459,  fig. 
8,  6.  Male  holotype  from  Santa  Ana,  Guatemala, 
in  BMNH.  Roewer,  1942:  868.  Bonnet,  1957:  2823. 

Araneus  uncatus: — Petrunkevitch,  1911:  321. 

Description.  Female  from  Ayutla,  San 
Marcos,  Guatemala.  Carapace  dark  brown 
with  light  region  surrounding  the  eyes  and 
extending  posteriorly  behind  the  lateral 
eyes.  Pair  of  darker  feather-shaped  patches 
anterior  of  thoracic  furrow,  touching  pos- 
teriorly to  create  a  U-shape  (see  Fig.  255). 
Dark  rings  on  distal  half  of  leg  articles,  an- 
terior side  of  patellae,  and  dark  markings 
on  dorsal,  proximal  end  of  tibia  I  and  II. 
Femur  I  with  four  macrosetae  on  anterior 
side;  one  on  anteroventral  side.  Dorsum  of 
abdomen  with  white  fleur-de-lis  folium; 
this  pattern  thinner  than  in  most  other 
species.  Fleur-de-lis  pattern  on  a  dark 
background,  outlined  on  either  side  by 
thin  white  stripe  (Fig.  255).  Dark  lateral 
band  follows  sides  of  abdomen  and 
stretches  up  over  anterior  dorsal  portion. 
Venter  with  two  white  spots  on  each  side 
of  spiracle;  wide,  white,  medial  longitudi- 
nal line  starts  anterior  to  the  colulus  and 
ends  posterior  to  the  epigynal  groove. 
Thin,  faint  lines  run  parallel  to  wide  me- 
dian one  anterior  to  white  spots.  White 
patch  separates  epigynum  and  dark  de- 
pression posterior  to  pedicel  (Fig.  256). 
Sternum  black  with  white  dewdrop- 
shaped  mark  at  posterior  end  (Fig.  256). 
Ratio  of  eye  diameters:  posterior  medians 
and  anterior  medians  1.0,  anterior  laterals 
1.3,  posterior  laterals  1.2.  Anterior  median 
eyes  separated  by  1.6  diameters,  posterior 
median  eyes  by  1.1,  anterior  median  eyes 
separated  froin  anterior  laterals  by  4  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.4  their  diameters.  To- 
tal length  7.8  mm.  Carapace  3.7  mm  long, 
2.9  wide.  First  femur  4  mm,  patella  and 
tibia  4.3,  metatarsus  3.8,  tarsus  1.4.  Sec- 
ond patella  and  tibia  3.6  mm,  third  2.3, 
fourth  3.2. 


Metepeira  •  Piel         77 


Male  from  Ayutla,  San  Marcos,  Guate- 
mala. Carapace  dirty  yellowish  brown  with 
lighter  mark  in  center  (Fig.  253).  Saine 
color  as  carapace  except  lighter  on  proxi- 
mal half  of  femora.  Femur  I  with  row  of 
nine  macrosetae  on  anterior  side;  five  on 
anteroventral  side.  Male  abdomen  similar 
to  female,  but  leaves  of  fleur-de-lis  pattern 
thinner  (Fig.  253).  Venter  with  a  shorter 
longitudinal  line,  as  compared  with  female 
(Fig.  254).  Sternum  with  posterior  white 
dewdrop  as  in  female,  and  additional  an- 
terior white  mark  near  labium  (Fig.  254). 
Ratio  of  eye  diameters:  posterior  medians 
and  anterior  medians  0.9,  anterior  laterals 
1.3,  posterior  laterals  1.3.  Anterior  median 
eyes  separated  by  1.5  diameters,  posterior 
median  eyes  by  1.0,  anterior  median  eyes 
separated  from  anterior  laterals  by  2  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.3  their  diameters.  To- 
tal length  4.5  mm.  Carapace  2.3  mm  long, 
1.8  wide.  First  femur  3.4  mm,  patella  and 
tibia  3.4,  metatarsus  3.2,  tarsus  1.1.  Sec- 
ond patella  and  tibia  2.7  mm,  third  1.4, 
fourth  2. 

Diagnosis.  The  generally  darker  pig- 
mentation (Fig.  255)  and  the  small  white 
mark  on  the  sternum  (Fig.  256)  separate 
M.  uncata  from  all  other  members  of  the 
M.  Ventura  species  group.  Like  M.  ventura 
(Fig.  231)  but  unlike  other  members  of 
the  M.  ventura  species  group,  the  open- 
ings to  the  epigynum  are  more  triangular 
to  oval  (Fig.  252),  rather  than  square  (Figs. 
259,  266).  Also  distinctive  are  the  sinuous 
black  lines  that  form  upside-down  U- 
shapes  inside  each  epigynal  opening  (Fig. 
252),  in  contrast  to  the  comma-shaped 
marks  in  M.  ventura  or  M.  chilapae  (Figs. 
231,  266).  Like  M.  chilapae  but  unlike  oth- 
er members  of  the  M.  ventura  group,  the 
median  apophysis  and  flagellae  of  M.  un- 
cata are  slimmer  and  of  uniform  thickness 
(Figs.  250,  264).  The  palp  of  M.  uncata 
differs  from  that  of  M.  chilapae  by  the 
more  basal  position  of  the  bend  in  the  em- 
bolus tip  (compare  Fig.  250  with  Fig.  264). 

Variation.  Average  body  length  of  five 
females  examined  8.2  mm,  range  7.5  to  8.8 


mm.  Average  body  length  of  four  males 
examined  5.6  mm,  range  4.5  to  6.4  mm. 

Natural  Historij.  Mature  specimens 
have  been  collected  in  August  (Fig.  329). 

Distribution.  Southwestern  Guatemala 
to  northern  Costa  Rica  (Map  12)  at  alti- 
tudes ranging  from  100  to  3,000  m. 

Records  Examined.  COSTA  RICA  Cordillera:  20 
km  N  Siquires,  10°9'N,  84°17'W,  15.viii.l980  (W. 
Eberhard,  MCZ).  GUATEMALA  El  Quiche:  Chichi- 
castenango,  14°56'N,  91°7'W,  6.viii.l947  (C.  &  P. 
Vaurie,  AMNH).  Huehuetenango:  Todos  Santos  Cu- 
chumatan,  15°31'N,  91°37'W,  16.viii.l979  (C.  E. 
Griswold,  CAS).  Quetzaltenango:  El  Baul,  1  km  S 
Quetzaltenango,  14°49'N,  91°31'W,  14.viii.l979  (T  C. 
Meikle  &  C.  E.  Griswold,  CAS);  Quetzaltenango, 
14°50'N,  91°31'W,  16.viii.l950  (C.  J.  &  M.  Good- 
night, AMNH).  San  Marcos:  Ayutla,  14°40'N,  92°9'W, 
19.viii.l947  (AMNH). 

33.  Metepeira  crassipes 
Chamberlin  and  Ivie 
Figures  257-263,  309;  Map  12 

Metepeira  josepha  Chamberlin  and  Ivie,  1942;  64,  fig. 
165,  9 .  Female  holotype  from  Kings  Mtn.  near 
Palo  Alto,  California  in  the  AMNH.  Synonymized 
by  Levi  (1977). 

Metepeira  crassipes  Chamberlin  and  Ivie,  1942:  66, 
figs.  171-173,  9,  S.  Male  holotype,  female,  male 
paratypes  from  Laguna  Beach,  California  in  the 
AMNH.  Levi,  1977:  202,  figs.  47-52,  9,6. 

Note.   As  first  reviser,  Levi  (1977)  preferred  to 
use  name  M.  crassipes. 

Description.  Female  from  Isla  San  Pe- 
dro Nolasco,  Sonora,  Mexico.  Carapace 
with  large  white  eye  region  and  lateral 
posterior  extensions  (Fig.  262).  Legs 
ringed  at  distal  ends  of  articles — though 
sometimes  only  lightly.  Femur  I  with  four 
macrosetae  on  anterior  side  aligned  in 
straight  row;  one  to  four  light  setae  on  an- 
teroventral side.  Dorsum  of  abdomen  with 
usual  Metepeira  folium  (Fig.  262);  venter 
wide,  with  long  white  median  line  set 
within  U-shaped  thinner  white  lines.  Pair 
of  white  spots  on  either  side  of  spiracle 
(Fig.  263).  Sternum  black  with  wide  me- 
dian white  line  widening  anteriorly  (Fig. 
263).  Ratio  of  eye  diameters:  posterior  me- 
dians and  anterior  medians  1.1,  anterior 
laterals  1.4,  posterior  laterals  1.3.  Anterior 
median  eyes  separated  by  1.4  diameters. 


78         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


posterior  median  eyes  by  0.8,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  3  diameters  of  anterior  lateral  eyes,  lat- 
eral eyes  separated  by  0.1  their  diameters. 
Total  length  5.7  mm.  Carapace  2.8  mm 
long,  2.1  wide.  First  femur  3.2  mm,  patella 
and  tibia  3.1,  metatarsus  2.8,  tarsus  1.  Sec- 
ond patella  and  tibia  2.7  mm,  third  1.6, 
fourth  2.5. 

Male  from  Winchester,  California.  Car- 
apace as  in  female  except  often  with  me- 
dian white  mark.  Femur  I  with  row  of  four 
macrosetae  on  anterior  side;  about  four  to 
eight  on  anteroventral  side.  Dorsum,  ven- 
ter as  in  female  (Fig.  260);  median  white 
line  on  sternum  more  often  broken  (Fig. 
261).  Ratio  of  eye  diameters:  posterior  me- 
dians and  anterior  medians  1.0,  anterior 
laterals  1.6,  posterior  laterals  1.4.  Anterior 
median  eyes  separated  by  1.2  diameters, 
posterior  median  eyes  by  0.6,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  2  diameters  of  anterior  lateral  eyes,  lat- 
eral eyes  separated  by  0.3  their  diameters. 
Total  length  4  mm.  Carapace  2.1  mm  long, 
1.5  wide.  First  femur  3.2  mm,  patella  and 
tibia  3.2,  metatarsus  3.2,  tarsus  1.1.  Sec- 
ond patella  and  tibia  2.5  mm,  third  1.3, 
fourth  2. 

Diagnosis.  Unlike  other  species  in  the 
M.  Ventura  species  group,  the  openings  to 
the  epigynum  of  M.  crassipes  resemble 
those  of  M.  chilapae  because  they  are 
shaped  like  squares  with  rounded  edges 
(compare  Fig.  259  with  Fig.  266).  How- 
ever, compared  to  M.  chilapae,  the  epigyn- 
al  openings  of  M.  crassipes  are  relatively 
smaller  and  the  scape  is  relatively  longer. 
The  embolus  of  M.  crassipes  is  distin- 
guished by  its  gentle  cui'vature  (Fig.  257), 
in  contrast  to  much  more  abi"upt  cui"vature 
seen  in  other  species  in  the  M.  ventura 
group  (e.g..  Fig.  264),  with  the  exception 
of  M.  revillagigeclo.  The  median  apophysis 
of  M.  crassipes  (Fig.  257)  is  slimmer  than 
that  of  M.  revillagigedo  (Fig.  236). 

Variation.  Average  body  length  of  four 
females  examined  6  mm,  range  5  to  6.5 
mm.  Average  body  length  of  two  males  ex- 
amined 4.4   mm,   range   4.1   to   4.7   mm. 


Sometimes  the  scape  is  greatly  swollen, 
adding  to  its  relative  width.  In  such  cases 
the  scape  can  cover  the  epigynal  openings, 
which  may  mislead  the  investigator  to  con- 
fuse it  for  Metepeira  arizonica. 

Natural  Histortj.  Levi  (1977)  reports 
that  males  in  the  U.S.  have  been  collected 
from  April  to  October,  primarily  in  Cali- 
fornia buckwheat  and  sage.  Mexican  re- 
cords expand  this  seasonality  to  include 
the  entire  year  (Fig.  309). 

Distribution.  From  northern  California 
in  the  U.S.  to  Baja  California  Sur,  Mexico 
(Levi,  1977,  map  1;  Map  12). 

Records  Examined.  MEXICO  Baja  Calif.  Norte: 
Santo  Tomas,  31°33'N,  116°24'W,  8.vii.l953  (W.  J.  & 
J.  W.  Gertsch,  AMNH),  12.xi.l976  (S.  C.  Williams  & 
K.  B.  Blair,  CAS).  Baja  Calf.  Sun  26  mi  S  Loreto, 
25°37'N,  111°17'W,  l.i.l977  (C.  E.  Griswold  &  L. 
Vincent,  CAS);  Isla  Magdalena,  Puerto  Magdalena, 
24°38'N,  112°9'W,  16.iii.l957  (R.  Zweifel,  AMNH); 
Isla  San  Francisco,  South  Side,  24°50'N,  110°35'W, 
19.V.1970  (S.  C.  Williams  &  V.  F.  Lee,  CAS);  Sierra 
San  Nicolas,  26°32'N,  111°36'W  [?]  (Eisen  &  Vaslit, 
MCZ).  Sonora:  Isla  San  Pedro  Nolasco,  27°58'N, 
111°25'W,  17.iv.l921  (G.  C.  Chamberlin,  MCZ);  Si- 
erra de  Alamos,  30°51'N,  112°2'W  [?],  19.i.l968  (V. 
Roth,  AMNH).  USA  California:  1  mi  NW  Winches- 
ter (Double  Butte),  in  web  between  Artemisia  cali- 
fornica  bushes,  33°43'N,  117°6'W,  4.xii.l976  (W. 
Icenogle,  MCZ). 

34.  Metepeira  chilapae 
Chamberlin  and  Ivie 
Figures  264-270,  333;  Map  13 

Metepeira  chilapae  Chamberlin  and  Ivie,  1936:  45, 
figs.  119-121,  S.  Male  holotype  from  Chilapa, 
Guerrero,  Mexico,  in  the  ZMB.  Roewer,  1942:  868. 

Metepeira  chilapica: — Bonnet,  1957:  2820.  Unjusti- 
fied eiuendation. 

Note.  Examination  of  Chamberlin  and  Ivies 
(1936)  and  Levi's  (personal  illustrations)  figures  of 
the  holotype  was  sufficient  to  identify  M.  chilapae 
accurately. 

Description.  Female  from  Cocoyoc, 
Morelos,  Mexico.  Brownish  black  cara- 
pace, paler  around  and  just  posterior  to 
lateral  eyes  (Fig.  269).  Distal  halves  of 
ventral  leg  articles  black,  elsewhere  yel- 
lowish. Femur  I  with  row  of  four  macro- 
setae  on  anterior  side;  zero  to  three  setae 
on  anteroventral  side.  Dorsal  folium  a 
white  fleur-de-lis  marking  set  on  golden- 


Metepeira  •  Piel        79 


257 


crassipes 
(33) 


260        261 


269 

chilapae 
(34) 


270 


276 


277 


272 


petatlan 
(35) 


Figures  257-263.  Metepeira  crassipes  Chamberlin  and  Ivie  (sp.  33  [257-261]  33°43'N,  117°6'W;  [262,263]  27°58'N,  1 1 1°25'W). 

257,  male  palpus,  mesal.  258,  epigynum,  posterior.  259,  epigynum,  ventral.  260,  male,  dorsal.  261,  male,  ventral.  262,  female, 

dorsal.  263,  female,  ventral. 

Figures  264-270.  Metepeira  cliilapae  Chamberlin  and  Ivie  (sp.  34  [264]  17°39'N,  99°22'W;  [265-270]  17°39'N,  99°22'W).  264, 

male  palpus,  mesal.  265,  epigynum,  posterior.  266,  epigynum,  ventral.  267,  male,  dorsal.  268,  male,  ventral.  269,  female,  dorsal. 

270,  female,  ventral. 

Figures  271-277.  Metepeira  petatlan  new  species  (sp.  35  [271,274,275]  17°14'N,  100°53'W;  [272,273,276,277]  17°17'32"N, 

101°2'40"W).  271,  male  palpus,  mesal.  272,  epigynum,  posterior.  273,  epigynum,  ventral.  274,  male,  dorsal.  275,  male,  ventral. 

276,  female,  dorsal.  277,  female,  ventral. 

Scale  bars:  dorsum  and  venter  figures  1.0  mm. 


80 


Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


brown  speckled  pattern  over  white.  Foli- 
um darkens  posteriorly  (Fig.  269).  Ante- 
rior shoulders  black.  Venter  black  with 
wide  median  white  line  and  pair  of  white 
spots  on  eidier  side  of  spiracle  (Fig.  270). 
Sternum  brownish  black  with  wide,  white 
line  widening  anteriorly  (Fig.  270).  Ratio 
of  eye  diameters:  posterior  medians  and 
anterior  medians  1.0,  anterior  laterals  1.1, 
posterior  laterals  1.1.  Anterior  median 
eyes  separated  by  1.3  diameters,  posterior 
median  eyes  by  0.8,  anterior  median  eyes 
separated  from  anterior  laterals  by  2.2  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.2  their  diameters.  To- 
tal length  5.8  mm.  Carapace  2.7  mm  long, 
2.3  wide.  First  femur  2.9  mm,  patella  and 
tibia  3,  metatarsus  2.6,  tarsus  1.  Second 
patella  and  tibia  2.4  mm,  third  1.6,  fourth 
2.3. 

Male  from  6  mi  northeast  of  Tixtla  de 
Guerrero,  Guerrero,  Mexico.  Carapace, 
dorsum,  venter,  sternum  darker  and  more 
contrasty  version  of  female  (Figs.  267, 
268).  Distal  halves  of  ventral  leg  articles 
black,  elsewhere  yellowish.  Femur  I  with 
row  of  four  macrosetae  on  anterior  side; 
three  on  an tero ventral  side.  Ratio  of  eye 
diameters:  posterior  medians  and  anterior 
medians  1.0,  anterior  laterals  1.3,  posterior 
laterals  1.3.  Anterior  median  eyes  separat- 
ed by  1.7  diameters,  posterior  median  eyes 
by  0.9,  anterior  median  eyes  separated 
from  anterior  laterals  by  1.9  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.3  their  diameters.  Total  length  3.5 
mm.  Carapace  1.8  mm  long,  1.4  wide. 
First  femur  2.3  mm,  patella  and  tibia  2.1, 
metatarsus  1.8,  tarsus  0.8.  Second  patella 
and  tibia  1.8  mm,  third  0.9,  fourth  1.4. 

Diagnosis.  The  openings  to  the  epigyn- 
um  resemble  those  of  M.  crassipes  be- 
cause they  are  shaped  like  squares  with 
rounded  edges  (compare  Fig.  266  with 
Fig.  259).  However,  the  dark,  comma- 
shaped  mark  inside  each  opening  resem- 
bles that  of  M.  Ventura  (compare  Fig.  266 
with  Fig.  231).  As  in  M.  ventura  (Fig.  229) 
but  unlike  all  other  species  in  the  M.  ven- 
tura  species   group,    the   embolus   tapers 


strongly  then  curves  sharply  up  and  inward 
right  at  the  tip  (Fig.  264).  The  bend  in  the 
embolus  in  other  M.  ventura  group  species 
is  less  pronounced  (Figs.  236,  257)  or  not 
so  near  the  tip  (Figs.  243,  250).  Unlike  M. 
ventura,  the  median  apophysis  in  M.  chi- 
lapae  is  slimmer  (Fig.  264)  and  the  sepa- 
ration between  the  embolus  and  the  basal 
embolic  apophysis  is  relatively  greater. 

Variation.  Average  body  length  of  eight 
females  examined  6.5  ixim,  range  5.7  to  8.3 
mm.  Average  body  length  of  three  males 
examined  2.9  mm,  range  2.3  to  3.5  mm. 

Natural  History.  Mature  specimens 
have  been  collected  in  July  through  Oc- 
tober (Fig.  333). 

Distribution.  Southern  Mexican  states 
between  Nayarit  and  Oaxaca;  elevations 
from  1,000  to  2,000  m  (Map  13). 

Records  Examined.  MEXICO  Guerrero:  6  mi  NE 
Tixtla  de  Guerrero,  17°39'N,  99°22'W,  16.vii.l984  (J. 
B.  Woolley,  ADC).  Morelos:  Cocoyoc,  18°52'N, 
98°59'W,  28.vii.1956  (W.  Gertsch  &  V.  Roth,  AMNH); 
Cuemavaca,  18°55'N,  99°15'W,  15.X.1944  (N.  L.  H. 
Krauss,  AMNH).  Nayarit:  30  mi  SE  Tepic,  21°12'N, 
104°33'W,  23.xi.1948  (E.  S.  Ross,  CAS);  5  mi  NW 
Tepic,  21°32'N,  104°57'W,  13.V.1963  (W.  J.  Gertsch 
&  W.  Ivie,  AMNH).  Oaxaca:  6  mi  NE  Mitla,  16°59'N, 
96°21'W,  20.viii.l985  (J.  Woolley  &  G.  Zolnerowich, 
ADC);  Huajuapan,  17°48'N,  97°46'W,  l.x.1946  (H. 
Wagner,  AMNH);  Oaxaca,  17°3'N,  96°43'W, 
17.vii.l955  (C.  &  R  Vaurie,  AMNH),  12.viii.l991  (W. 
H.  Piel  &  G.  S.  Bodner,  MCZ);  San  Bait.  Chichica- 
pan,  16°45'N,  96°29'W,  4.viii.l991  (W.  H.  Piel  &  G. 
S.  Bodner,  MCZ). 

Metepeira  minima  Group 

Female  spiders  in  the  M.  minima  group 
(Metepeira  petatlan,  Metepeira  uninima, 
Metepeira  pacifica,  Metepeira  jamaicensis) 
have  a  thin  scape  with  epigynal  openings 
that  are  shaped  from  longitudinal  slits 
(Figs.  273,  280,  281)  to  ovals  that  are  lon- 
ger than  wide  (Figs.  288,  295).  In  males, 
the  flagellae  on  the  median  apophysis  are 
set  off  on  a  distinctly  narrower  stalk  (Figs. 
271,  278,  286,  293). 

35.  Metepeira  petatlan  new  species 
Figures  271-277,  334;  IVIap  10 

Holotype.   Male  from  50  km  southeast  of  Petatlan, 
Guerrero,   Mexico,   14.viii.l984,  J.   B.  Woolley,  in 


Metepeira  •  Piel 


81 


MCZ.  The  specific  name  is  a  noun  in  apposition 
after  the  locality. 

Description.  Female  paratype  from  Pa- 
panoa,  Guerrero.  Light  region  around  pos- 
terior eye  row;  carapace  darkens  posteri- 
orly, then  lightens  under  overhang  of  ab- 
domen (Fig.  276).  Slight  rings  on  patella 
and  tibia.  Femur  I  with  three  macrosetae 
on  anterior  side,  none  on  anteroventral 
side.  Dorsum  of  abdoinen  with  typical  fo- 
lium, except  that  white  oak  leaf  pattern 
narrower  than  most  species  and  set  on  a 
narrow,  remarkably  darker  brownish  dove- 
tail (Fig.  276).  Dark  lateral  band  wraps 
around  abdomen  and  stretches  up  over  an- 
terior dorsal  portion.  Venter  with  two  spots 
on  either  side  of  spiracle  and  slight  indi- 
cation of  a  transverse  bar.  Wide,  white  lon- 
gitudinal mark  ends  at  the  epigynal  groove 
(Fig.  277).  Sternum  with  wide,  white  lon- 
gitudinal mark  v^dening  anteriorly  (Fig. 
277).  Ratio  of  eye  diameters:  posterior  me- 
dians and  anterior  medians  1.0,  anterior 
laterals  1.3,  posterior  laterals  1.2.  Anterior 
median  eyes  separated  by  1.1  diameters, 
posterior  median  eyes  by  0.7,  anterior  me- 
dian eyes  separated  from  anterior  laterals 
by  1  diameter  of  anterior  lateral  eyes,  lat- 
eral eyes  aliTiost  touching.  Total  length  5 
mm.  Carapace  2.2  mm  long,  1.7  wide. 
First  femur  2.3  mm,  patella  and  tibia  2.5, 
metatarsus  2,  tarsus  0.9.  Second  patella 
and  tibia  2  mm,  third  1.3,  fourth  1.9. 

Male  holotype.  Carapace  uniform 
brown  with  a  lighter  median  streak  just  an- 
terior to  thoracic  depression  (Fig.  274). 
Weakly  ringed  on  tibia;  distal  half  of  fem- 
ora dark.  Feinur  I  with  two  macrosetae  on 
anterior  side.  Abdomen  is  a  lighter  version 
of  the  female  (Fig.  274).  Venter  with  a 
much  reduced  and  shorter  longitudinal 
line,  compared  to  the  female  (Fig.  275). 
Sternum  as  in  female.  Ratio  of  eye  diam- 
eters: posterior  medians  and  anterior  me- 
dians 1.0,  anterior  laterals  1.4,  posterior 
laterals  1.4.  Anterior  median  eyes  separat- 
ed by  1.1  diameters,  posterior  median  eyes 
by  0.7,  anterior  median  eyes  separated 
from  anterior  laterals  by  1.1  diameters  of 


anterior  lateral  eyes,  lateral  eyes  separated 
by  0.2  their  diameters.  Total  length  2  mm. 
Carapace  1  mm  long,  0.7  wide.  First  fe- 
mur 1.2  inm,  patella  and  tibia  1.0,  meta- 
tarsus 0.8,  tarsus  0.5.  Second  patella  and 
tibia  0.9  mm,  third  0.5,  fourth  0.8. 

Diagnosis.  The  female's  dorsum  can  be 
distinguished  from  other  species  by  the 
unusually  dark  brown  color  of  the  dovetail- 
shaped  mark  (Fig.  276).  Both  this  mark 
and  the  white  oak  leaf  pattern  have  nar- 
rower margins  that  are  more  parallel  and 
not  as  wedge-shaped  as  they  are  in  other 
species  (Fig.  276).  The  epigynum  resem- 
bles that  of  M.  uninima  by  the  narrow 
scape  and  the  parallel  slitlike  openings  on 
either  side.  These  openings  differ  from 
those  of  M.  minima  because  they  have 
conspicuous  sclerotized  spheres  just  ante- 
rior to  the  lateral  edge  of  the  slits  (Fig. 
273)  but  which  are  much  farther  away  in 
M.  minima  (Fig.  280).  The  male  is  unusu- 
ally small  and  its  palp  is  unique  and  easily 
distinguished  from  other  species.  The  lon- 
ger flagellum  on  the  median  apophysis  is 
needlelike  over  its  entire  length,  not  grad- 
ually tapering  (compare  Fig.  271  with  Fig. 
278).  In  addition,  the  long  needlelike  fla- 
gelluni  has  a  sharper  elbow.  At  the  elbow 
it  projects  away  from  the  palp  for  a  short 
distance  and  then  strongly  curves  around 
the  palp  (Fig.  271). 

Natural  History.  The  female  paratype 
was  found  at  eye  level  on  a  hot,  dry,  wood- 
ed hillside  overlooking  the  Pacific  Ocean 
at  an  altitude  of  about  200  m.  Her  web 
differed  considerably  from  the  usual  Me- 
tepeira web:  instead  of  an  elaborate  barrier 
web,  it  consisted  of  a  much  reduced  Y- 
shaped  structure  with  a  curled  leaf  in  the 
center.  The  leaf  served  as  a  retreat  for  the 
spider  and  protected  and  hid  three  egg 
sacs.  The  retreat  was  suspended  very  near 
the  hub,  as  opposed  to  farther  away  and 
above  it,  as  found  in  most  other  species. 
Mature  specimens  have  been  collected  in 
August  through  October  (Fig.  334). 

Distribution.  Western  Mexico:  Guerrero 
and  Sinaloa  (Map  10). 


82         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


Records  Examined.  MEXICO  Guerrero:  32  mi  SE 
Petatlan,  17°14'N,  100°53'W,  14.viii.l984  (J.  B.  Wool- 
ley,  MCZ);  Microondas  Tamarindos,  S.  Papanoa, 
17°17'32"N,  101°2'40"W,  18.x. 1994  (W.  H.  Piel, 
MCZ).  Sinaloa:  2  mi  S  Elota,  23°55'N,  106°48'W, 
ll.Lx.1966  (Jean  &  Wilton  Ivie,  AMNH). 

36.  Metepeira  minima  Gertsch 
Figures  278-285,  328;  Map  15 

Metepeira  minima  Gertsch,  1936:  10,  fig.  31,  6 .  Male 
hoiotype  from  Edinburg,  Texas,  in  the  AMNH,  ex- 
amined. Roewer,  1942:  869.  Chamberlin  and  Ivie, 
1942:  67,  fig.  174,  9.  Levi,  1977:  206,  fig.  70-73, 
76-77,  9 ;  74-75,  3 .  Bonnet,  1957:  2822.  Brignoli, 
1983:  275. 

Description.  Female  from  Celestun,  Yu- 
catan, Mexico.  Carapace  dark  brown  with 
light  region  surrounding  the  eyes,  some- 
times extending  posteriorly  behind  the  lat- 
eral eyes.  Faint  pair  of  darker  feather- 
shaped  patches  in  center  of  carapace  (Fig. 
284).  Legs  same  color  as  carapace;  darker 
on  distal  ends  of  articles.  Femur  I  with 
row  of  four  macrosetae  on  anterior  side; 
rarely  any  on  anteroventral  side.  Dorsum 
of  abdomen  with  usual  Metepeira  folium 
(Fig.  284);  venter  wide,  with  long  white 
median  line  surrounded  by  faint,  thin, 
white  U-shaped  line  (Fig.  285).  Pair  of 
large  white  spots  on  either  side  of  spiracle. 
Sternum  black  with  wide,  white,  uneven 
median  line  (Fig.  285).  Ratio  of  eye  di- 
ameters: posterior  medians  and  anterior 
medians  1.0,  anterior  laterals  1.2,  posterior 
laterals  1.1.  Anterior  median  eyes  separat- 
ed by  1.2  diameters,  posterior  median  eyes 
by  0.9,  anterior  median  eyes  separated 
from  anterior  laterals  by  1.9  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.2  their  diameters.  Total  length  6.5 
mm.  Carapace  2.7  mm  long,  2  wide.  First 
femur  2.8  mm,  patella  and  tibia  2.8,  meta- 
tarsus 2.5,  tarsus  0.9.  Second  patella  and 
tibia  2.4  mm,  tliird  1.5,  fourth  2. 

Male  from  Celestun,  Yucatan,  Mexico. 
Carapace  dark  with  light  mark  anterior  to 
thoracic  furrow  (Fig.  282).  Femur  I  with 
row  of  four  macrosetae  on  anterior  side; 
row  of  two  to  five  macrosetae  on  anter- 
oventral side.  Distal  two-thirds  same  color 
as  carapace,  proximal  third  white.  Folium 


of  abdomen  as  in  female,  except  posterior 
half  darker  than  anterior  half  (Fig.  282). 
Venter  of  abdomen  and  sternum  as  in  fe- 
male, except  median  white  line  of  sternum 
often  broken  (Fig.  283).  Ratio  of  eye  di- 
ameters: posterior  medians  and  anterior 
medians  1.1,  anterior  laterals  1.4,  posterior 
laterals  1.1.  Anterior  median  eyes  separat- 
ed by  1.2  diameters,  posterior  median  eyes 
by  0.8,  anterior  median  eyes  separated 
from  anterior  laterals  by  1.5  diameters  of 
anterior  lateral  eyes,  lateral  eyes  separated 
by  0.1  their  diameters.  Total  length  3  mm. 
Carapace  1.5  mm  long,  1.2  wide.  First  fe- 
mur 2  mm,  patella  and  tibia  1.9,  metatar- 
sus 1.8,  tarsus  0.8.  Second  patella  and  tibia 
1.6  mm,  third  0.8,  fourth  1.3. 

Diagnosis.  Unlike  other  members  of  the 
M.  niininia  species  group,  the  openings  to 
the  epigynum  of  M.  minima  are  narrow 
slits  around  a  scape  of  variable  size  (Figs. 
280,  281).  The  openings  of  M.  petatlan  are 
only  slightly  wider  than  M.  minima,  but 
the  internal  darker  sclerotized  spheres  in 
M.  minima  sit  much  farther  apart  (com- 
pare Fig.  280  with  273).  Unlike  M.  pacifica 
(Fig.  286)  and  M.  jamaicensis  (Fig.  293), 
the  bent  embolus  on  M.  minima  tapers 
strongly  and  is  therefore  not  like  a  needle 
(Fig.  278).  Unlike  M.  petatlan,  the  longer 
flagellum  on  M.  minima  tapers  (Fig.  278) 
and  is  not  thin  over  its  entire  length  (Fig. 
271). 

Variation.  Average  body  length  of  13  fe- 
males examined  5.4  mm,  range  3.7  to  6.5 
mm.  Average  body  length  of  17  males  ex- 
amined 3.4  mm,  range  2.5  to  4.5  mm.  The 
epigyna  of  females  from  the  Yucatan  pen- 
insula differ  noticeably  from  all  others,  but 
the  males  hardly  show  any  distinguishing 
features.  These  differences  can  be  seen  in 
the  much  wider  scape  and  the  greater  sep- 
aration between  openings  in  a  specimen 
from  the  Yucatan  (Fig.  281),  compared  to 
a  specimen  from  Tamaulipas  (Fig.  280)  or 
one  from  Texas  (Levi,  1977:  209,  fig.  71). 
In  addition,  the  posterior  view  of  the  epi- 
gynum from  a  Yucatan  specimen  shows 
the  lobes  converging  to  form  a  V-shape 
(Fig.  279),  whereas  females  outside  of  the 


Metepeira  •  Piel 


83 


278 


minima 
(36) 


s  291 


292 


289  ^     290 


pacifica 

(37) 


298 


297 


jamaicensis 
(38) 


Figures  278-285.  Metepeira  minima  Gertsch  (sp,  36  [278,279,281-285]  20°56'N,  90°21'W;  [280]  22°30'N,  99°4'W).  278,  male 

palpus,  mesal.  279,  epigynum,  posterior.  280,281,  epigynum,  ventral.  282,  male,  dorsal.  283,  male,  ventral.  284,  female,  dorsal. 

285,  female,  ventral. 

Figures  286-292.  Metepeira  pacifica  new  species  (sp.  37;  10°27'N,  85°9'W).  286,  male  palpus,  mesal.  287,  epigynum,  posterior. 

288,  epigynum,  ventral.  289,  male,  dorsal.  290,  male,  ventral.  291,  female,  dorsal.  292,  female,  ventral. 

Figures  293-299.  Metepeira  jamaicensis  Archer  (sp.  38;  18°17'N,  76°48'W).  293,  male  palpus,  mesal.  294,  epigynum,  posterior. 

295,  epigynum,  ventral.  296,  male,  dorsal.  297,  male,  ventral.  298,  female,  dorsal.  299,  female,  ventral. 

Scale  bars:  dorsum  and  venter  figures  1 .0  mm. 


84         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


Yucatan  show  largely  parallel  lobes  (Levi, 
1977:  209,  fig.  71). 

Natural  History.  Adults  have  been  ob- 
served on  tree  trunks  and  on  bushes  at  and 
above  150  cm.  They  can  be  found  in  shad- 
ed areas,  which  is  unusual  for  Mexican 
Metepeira.  Most  mature  specimens  have 
been  collected  in  May  through  September 
(Fig.  328).  Elevations  range  from  sea  level 
to  just  under  2,000  m. 

Distribution.  Southern  Texas  to  Hon- 
duras (Map  15). 

Records  Examined.  HONDUFIAS  Copdn:  14°50'N, 
89°9'W,  7.iii.l939  (AMNH);  Ruinas,  14°50'N, 
89°9'W,  7.iii.l939  (AMNH).  MEXICO  Campeche: 
Becan,  18°33'N,  89°30'W,  31.vii.l991  (W.  H.  Piel  & 
G.  S.  Bodner,  MCZ);  Campeche,  19°51'N,  90°32'W, 
14.vii.l948  (C.  J.  Goodnight,  AMNH);  Champoton 
beach,  19°21'N,  90°43'W,  22.vii.1991  (W.  H.  Piel  & 
G.  S.  Bodner,  MCZ);  Seybaplaya,  19°39'N,  90°40'W, 
2.vlii.l949  (C.  J.  Goodnight,  AMNH).  Chiapas:  16  mi 
W  Cintalpa  on  rt  190.  16°36'N,  93°51'W,  15.vi.l982 
(F.  Coyle,  MCZ).  Nuevo  Leon:  20  km  E  Montemo- 
relos,  Gamine  Q.  Rayones,  25°16'N,  99°41'W, 
22.vi.1981  (L.  Stange,  FSCA);  Los  Cristales,  25°33'N, 
100°12'W,  15.viii.l972  (A.  F.  Archer,  AMNH);  Villa 
de  Santiago,  Hacienda  Vista  Hermosa,  25°25'N, 
100°9'W,  19.vi.l940  (H.  Hoogstraal,  MCZ);  Villa  San- 
tiago, Horsetail  Falls,  25°25'N,  100°9'W,  19.vi.l940 
(H.  Hoogstraal,  MCZ).  San  Luis  Potosi:  20  km  W  Cd. 
Valles,  21°58'N,  99°11'W,  18.iii.l972  (J.  A.  L.  Cooke, 
AMNH);  Cd.  Valles,  21°59'N,  99°1'W,  6.vii.l940  (R 
Ran,  MCZ),  19.vli.l956  (W.  J.  Gertsch  &  V.  Roth, 
AMNH),  15.vii.l959  (L.  Steude,  AMNH);  Cd.  Valles, 
Hotel  Covadonga,  21°59'N,  99°1'W  (L.  Steude, 
AMNH);  Huichichuyan,  21°25'N,  98°55'W,  19.V.1952 
(M.  Cazier,  W.  Gertsch,  &  R.  Schrammel,  AMNH); 
Medina,  24°1'N,  100°24'W,  9.Lx.l956  (A.  F.  Archer, 
AMNH);  N  section  of  San  Luis  Potosi,  22°13'N, 
100°58'W,  8.ix.l956  (A.  F.  Archer,  AMNH);  Venado 
Arroyo,  22°56'N,  101°5'W  [?],  27.vii.1934  (MCZ).  So- 
nora:  8  mi  W  Alamos,  29°13'N,  110°10'W, 
23.viii.1965  (W.  J.  Gertsch  &  R.  Hastings,  AMNH). 
Tamaulipas:  60  km  N  Cd  Valles,  22°30'N,  99°4'W, 
10.viii.l991  (W.  H.  Piel  &  G.  S.  Bodner,  MCZ);  Cd. 
Victoria,  23°44'N,  99°8'W,  17.V.1952  (M.  Cazier,  W. 
Gertsch,  &  R.  Schrammel,  AMNH);  Laredo  road 
near  Cd.  Victoria,  23°44'N,  99°8'W,  20.viii.l947  (C. 
J.  &  M.  Goodnight,  AMNH);  rlOl  26km  S  Tula, 
22°49'N,  99°55'W,  8.Lx.l991  (W.  H.  Piel  &  G.  S.  Bod- 
ner, MCZ);  Sisal,  15  mi  S  Cd.  Victoria,  23°38'N, 
99°12'W,  22.vii.1966  (Jean  &  Wilton  Ivie,  AMNH). 
Veracruz:  15  mi  E  Panuco,  22°10'N,  98°3'W, 
29.xi.1941  (A.  M.  &  L.  I.  Davis,  AMNH);  Plan  del 
Rio,  19°6'N,  96°6'W  [?],  26.vii.1956  (W.  J.  Gertsch  & 
V.  Roth,  AMNH).  Yucatan:  20  km  E  Valladolid, 
20°41'N,  88°2'W,  26.vii.1991   (W.   H.  Piel  &  G.   S. 


Bodner,  MCZ);  3  km  S  San  Felipe,  21°32'N, 
88°14'W,  25.vii.1991  (W.  H.  Piel  &  G.  S.  Bodner, 
MCZ);  5  mi  E  Sisal  salt  flat,  21°9'N,  90°5'W,  9.i.l984 
(V.  &  B.  Roth,  CAS);  Balankanche  Cave,  2  km  E  Chi- 
chen  Itza,  20°40'N,  88°33'W,  19.vii.l983  (W.  Mad- 
dison,  MCZ);  beach  north  of  Celestun,  20°56'N, 
90°21'W,  24.vii.1991  (W.  H.  Piel  &  G.  S.  Bodner, 
MCZ);  Chichen  Itza,  20°40'N,  88°34'W  (C.  J.  Good- 
night, AMNH),  15.vii.l981  (C.  Gold,  CAS);  Cordil- 
leria  Mayapan,  20°28'N,  89°11'W,  6.Lx.l952  (J.  &  D. 
Pallister,  AMNH);  Uxmal,  20°22'N,  89°46'W, 
7.ix.l970  (A.  F.  Archer,  AMNH),  23.vii.1991  (W.  H. 
Piel  &  G.  S.  Bodner,  MCZ).  USA  Texas:  29  mi  S 
Sarita,  26°47'N,  97°47'W,  14..xi.l958  (A.  Brady, 
MCZ);  1  mi  S  Pharr,  26°10'N,  98°11'W,  14.xi.l958 
(A.  Brady,  MCZ);  1  mi  S  Pharr  on  U.S.  HW  281, 
26°10'N,  98°11'W,  14..xi.l95S  (A.  Brady,  MCZ). 

37.  Metepeira  pacifica  new  species 
Figures  286-292,  302;  Map  10 

Holotijpe.  Male  from  La  Pacifica,  Guanacaste,  Costa 
Rica,  I.ii.l975-2.iii.l975,  R.  E.  Coville,  in  MCZ. 
The  specific  name  is  a  noun  in  apposition  after  the 
locality. 

Description.  Female  paratype  from  La 
Pacifica,  Guanacaste,  Costa  Rica.  Carapace 
tan,  lighter  around  eyes.  Legs  white.  Fe- 
mur I  with  row  of  three  macrosetae  on  an- 
terior side;  none  on  anteroventral  side. 
Dorsum  of  abdomen  white  with  faint,  in- 
distinct folium,  darker  distally  (Fig.  291). 
Gravid  females  often  slightly  marbled. 
Venter  wide,  with  long  white  median  line 
surrounded  by  faint,  thin,  white  U-shaped 
line  on  black.  Pair  of  large  white  spots  on 
each  side  of  spiracle  (Fig.  292).  Sternum 
black  with  wide,  white,  uneven  median 
line  (Fig.  292).  Ratio  of  eye  diameters: 
posterior  medians  and  anterior  medians 
1.0,  anterior  laterals  1.2,  posterior  laterals 
1.2.  Anterior  median  eyes  separated  by  1.2 
diameters,  posterior  median  eyes  by  0.9, 
anterior  median  eyes  separated  from  an- 
terior laterals  by  1.8  diameters  of  anterior 
lateral  eyes,  lateral  eyes  separated  by  0.2 
their  diameters.  Total  length  4.5  mm.  Car- 
apace 2.2  mm  long,  1.7  wide.  First  femur 
2.3  mm,  patella  and  tibia  2.4,  metatarsus 
1.9,  tarsus  0.8.  Second  patella  and  tibia  2 
mm,  third  1.2,  fourth  1.8. 

Male  holotype.  Carapace,  legs,  abdo- 
men as  in  female,  though  often  darker 
(Figs.  289,  290).  Femur  I  with  row  of  four 


Metepeira  •  Piel 


85 


300  Dec 

Nov 


301 


gressa,  n  =  26 


Sep 


Dec 

Jan 

Nov  ^ 

< 

z 

'^Mar 

V 

-r* 

W"" 

Aug^ 

\ 

Julyl 

June 

revillagigedo,  n  =  2 


304  Dec 

Nov 


Apr  Sep 


galatheae,  n  =  304 


tarapaca,  n  =  23 


July 

rectangula,  n  =  1 1 


desenderi,  n  =  46 


308  De': 

Nov 


calamuchita,  n  =  15 


Nov 

■\ 

\"^ 

OaA 

1 

Am 

'"V 

\y 

V^ 

Aug 

y^  ^ 

May 

July 

rjune 

crassipes,  n 


May 
July     I     June 

nigriventris,  n  =  44 


Aug  NC^    ^     jP\  ^^y 

July    I     June 

compsa,  n  =  279 


datona,  n  =  66 


jamaicensis,  n  =  24 


glomerabilis,  n  =  33 


cajabamba,  n  =  1 1 


Aug  "V^        1      Js/  May 
July  June 

roraima,  n  =  11 


maya,  n  =  1 3 


Aug  ^yl  ■"      \^  May 

July  June 

inca,  n  =  29 


Aug  ^^     ^1        J^  May 
July     '      June 

arizonica,  n  =  45 


Figures  300-319.  Circular  liistograms  depicting  relative  seasonal  abundance  of  collecting  events  for  mature  spiders.  300-308. 
Primarily  collected  during  the  northern  hemisphere  winter  and  spring  seasons.  300,  Metepeira  gressa;  301,  Metepeira  revillagi- 
gedo; 302,  Metepeira  pacifica;  303,  Metepeira  karkii;  304,  Metepeira  galatheae;  305,  Metepeira  tarapaca;  306,  Metepeira  rec- 
tangula; 307,  Metepeira  desenderi;  308,  Metepeira  calamuchita.  309-314.  Generally  collected  throughout.  309,  Metepeira  cras- 
sipes; 310,  Metepeira  nigriventris;  311,  Metepeira  compsa;  312,  Metepeira  datona;  313,  Metepeira  jamaicensis;  314,  Metepeira 
glomerabilis.  315-319.  Primarily  collected  during  the  Northern  Hemisphere  summer  season.  315,  Metepeira  cajabamba;  316, 
Metepeira  roraima;  317,  Metepeira  maya;  318,  Metepeira  inca;  319,  Metepeira  arizonica. 


86 


Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


macrosetae  on  anterior  side;  two  to  three 
on  antero ventral  side.  Median  white  line 
on  sternum  often  broken  (Fig.  290).  Ratio 
of  eye  diameters:  posterior  medians  and 
anterior  medians  1.0,  anterior  laterals  1.6, 
posterior  laterals  1.6.  Anterior  median 
eyes  separated  by  1.1  diameters,  posterior 
median  eyes  by  0.6,  anterior  median  eyes 
separated  from  anterior  laterals  by  1.4  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  0.3  their  diameters.  To- 
tal length  3.4  mm.  Carapace  1.7  mm  long, 
1.2  wide.  First  femur  2.2  mm,  patella  and 
tibia  2,  metatarsus  1.7,  tarsus  0.8.  Second 
patella  and  tibia  1.6  mm,  third  0.9,  fourth 
1.3. 

Diagnosis.  Unlike  other  members  of  the 
M.  minima  species  group,  M.  pacifica  and 
M.  jamaicensis  share  very  light  pigmenta- 
tion, and  the  embolus  in  both  species  is 
needle-shaped  (Figs.  286,  289,  291,  293, 
296,  298).  Unlike  M.  jamaicensis,  the  fla- 
gellae  on  the  median  apophysis  of  M.  pa- 
cifica are  set  off  on  a  short,  wide  stalk  (Fig. 
286)  rather  than  a  long,  thin  one  (Fig. 
293).  A  ventral  view  of  the  epigynum  in 
M.  pacifica  shows  a  ridge  under  the  scape 
that  is  more  V-shaped  (Fig.  288),  com- 
pared to  a  straighter  line  (Fig.  295). 

Variation.  Average  body  length  of  three 
females  examined  5.3  mm,  range  4.5  to  5.8 
mm.  Average  body  length  of  five  males  ex- 
amined 3  mm,  range  2.8  to  3.4  mm.  Most 
specimens  appear  whitish,  but  the  loss  of 
pigment  is  variable,  especially  among 
males. 

Natural  History.  Specimens  have  been 
collected  from  the  wasp  nests  of  Trtjpar- 
gilum  nitidum,  T.  tenoctitlan,  and  T.  hen- 
soni.  In  Costa  Rica,  mature  specimens 
have  been  collected  in  November  through 
February;  Honduras  and  Nicaragua  in  July 
(Fig.  302).  Altitudes  range  from  100  to 
2,600  m. 

Distribution.  Costa  Rica,  Honduras,  and 
Nicaragua  (Map  10). 

Records  Examined.  COSTA  RICA  Guanacaste:  4 
km  NW  Canas,  La  Pacifica,  prey  of  Trypargilum  ben- 
soni,  10°27'N,  85°9'W,  29.1.1975  (R.  E.  Coville, 
MCZ);  4  km  NW  Canas,  La  Pacifica,  prey  of  Trypar- 


gilum nitidum,  10°27'N,  85°9'W,  I.ii.l975-2.iii.l975 
(R.  E.  Coville,  MCZ);  4  km  NW  Caiias,  La  Pacifica, 
prey  of  Trypargilum  tenoctitlan,  10°27'N,  85°9'W 
l.ii.l975  (R.  E.  Coville,  MCZ);  4  km  NW  Canas,  La 
Pacifica,  wasp  collected,  10°27'N,  85°9'W,  16.ii.l975 
(R.  E.  Coville,  MCZ);  Bagaces,  Palo  Verde,  10°21'N, 
85°21'W  19.1.1978  (W.  Eberhard,  MCZ);  ca.  Canas, 
10°25'N,  85°7'W,  15.xi.l982  (W  Eberhard,  MCZ); 
Canas,  10°25'N,  85°7'W  15.xl.l982  (W  Eberhard, 
MCZ).  Puntarena.s:  Elnca  San  Miento-Sialas,  10°9'N, 
84°54'W,  5.11.1976  (Roth  &  Schroepfer,  AMNH).  San 
Jose  [?]:  Santa  Maria,  9°39'N,  83°58'W,  15.1.1930 
(Dodge,  MCZ).  HONDURAS  Tegucigalpa:  Teguci- 
galpa, 14°6'N,  87°13'W,  l.vii.l948  (Clarke,  AMNH). 
NICARAGUA  Managua:  Laguna  de  Jiloa,  SW  Ma- 
nagua, campsite,  12°13'N,  86°19'W  8.vil.l970  (S. 
Riechert,  SR). 

38.  Metepeira  jamaicensis  Archer 
Figures  293-299,  313;  Map  15 

Metepeira  jamaicensis  Archer,  1958:  16,  fig.  33,  9 . 
Female  holotype  from  Port  Henderson,  St.  Cath- 
erine Parish,  Jamaica,  in  the  AMNH,  examined. 

Metepeira  rmnima: — Levi,  1977:  206,  208.  BrignoH, 
1983:  275.  Erroneous  synonymy. 

Description.  Female  from  Saint  Mary's 
Parish,  Strawberry  Fields  near  Robin  s  Bay 
and  Green  Castle,  Jamaica.  Carapace  dirty 
brown,  white  around  eyes,  central  w^hite 
wedge  (Fig.  298).  Legs  whitish  yellow; 
slight  rings  on  legs  II  and  III.  Femur  I 
with  row  of  three  to  four  macrosetae  on 
anterior  side;  none  or  only  a  few  very  fine 
setae  on  anteroventral  side.  Dorsal  folium 
lighter  than  in  most  species;  fleur-de-lis 
white  on  speckled  light  gray  (Fig.  298). 
Venter  brownish  gray  with  white  margins. 
Wide  median  white  line  with  pair  of  large 
white  spots  on  either  side  of  spiracle  (Fig. 
299).  Sternum  brownish  black  with  wide, 
white  line  widening  anteriorly,  sometimes 
broken  in  center  (Fig.  299).  Ratio  of  eye 
diameters:  posterior  medians  and  anterior 
medians  1.0,  anterior  laterals  1.2,  posterior 
laterals  1.1.  Anterior  median  eyes  separat- 
ed by  1.4  diameters,  posterior  median  eyes 
by  0.7,  anterior  median  eyes  separated 
from  anterior  laterals  by  2  diameters  of  an- 
terior lateral  eyes,  lateral  eyes  separated  by 
0.2  their  diameters.  Total  length  5.1  mm. 
Carapace  2.1  mm  long,  1.5  wide.  First  fe- 
mur 2.1  mm,  patella  and  tibia  2.2,  meta- 


Metepeira  •  Piel 


87 


Aug  NC^    ^t         \^   May 
July    I     June 

Ventura,  n  =  35 


Aug  N4/     1        J\     May 
July     I      June 

grandiosa  grandiosa,  n  =  4 


22 

Dec  J 

Jan 

Nov  , 

Feb 

OctN 

-^Mar 

Sepi^ 

J*, 

Aug 

July 

June 

•'^^May 

gosoga,  n  =  2 


Aug     7^^  J\     May 

July     I      June 

incrassata,  n  =  33 


Aug  N/       W\  \;     ^^"^ 

July     I      June 

grandiosa  alpina,  n  =  5 


326 

Dec 

Jan 

Nov 

7^ 

Feb 

Oct /^ 

^ 

Am^ 

Sep\ 

J^ 

Jap. 

Aug 

\ 

May 

July 

June 

327 

Dec 

Jan 

Nov 

7^ 

Feb 

Oct  A 

■V 

"^Mar 

Sepi^ 

^ 

/  Apr 

Aug^ 

\ 

May 

July 

June 

celestun,  n  =  8 


vigilax,  n  =  10 


28 

Dec  J 

Jan 

NOV/ 

\ 

Feb 

Oct?^ 

f^ 

\  Mar 

Sep\> 

V 

\    /  ^^' 

Aug^ 

/r 

May 

juijn 

June 

329 


Sep 


Dec 
Nov  J\ 

Jan 

^\^Mar 

^ 

J*" 

AugNc^^ 

\ 

May 

J^ 

\lune 

30 

Nov   X 

Dec 

Jan 

Feb 

Oc.?^ 

^ 

-^Mar 

Sep^ 

K 

-^Apr 

AugV 

;4 

May 

July 

June 

minima,  n  =  39 


uncata,  n  =  6 


pimungan,  n  =  6 


Comanche,  n  =  2 


332 

Dec 

Jan 

Nov 

y^ 

^Feb 

OoN 

V 

-A  Mar 

.pi^ 

^ 

J  Apr 

Aug 

\ 

May 

Julyl 

June 

333 

Dec  J 

Jan 

Nov 

\ 

Feb 

Oct /^ 

j^ 

•^A  Mar 

Sep\ 

\t 

Ja. 

Aug 

YJ 

May 

July 

Fjune 

34 

Nov   X 

Dec 

Jan 

Feb 

o„/V 

J^ 

'^A  Mar 

sS^ 

V 

■\    /  ^P' 

Aug^ 

^ 

May 

July 

Ijune 

335 

Dec 

Jan 

Nov 

\ 

^Feb 

OctN 

1^ 

-^Mar 

'"U 

|L 

J  Apr 

Aug 

^ 

/ 

May 

jiiiTl 

rTune 

lacandon,  n  =  10 


chilapae,  n  =  15 


petatlan,  n  =  3 


spinipes,  n  =  1 16 


336 

Dec  J 

Jan 

Nov  , 

Feb 

Oct  r^ 

•^A  Mar 

Sep  y     . 

V4 

^Apr 

Aug 

July 

June 

May 

atascadero,  n  =  1 7 


triangularis,  n  =  19 


Figures  320-337.  Circular  histograms  depicting  relative  seasonal  abundance  of  collecting  events  for  mature  spiders.  320-331, 
Primarily  collected  during  the  northern  hemisphere  summer  season.  320,  Metepeira  ventura;  321,  Metepeira  grandiosa  gradiosa; 
322,  Metepeira  gosoga;  323,  Metepeira  ventura;  324,  Metepeira  olmec;  325,  Metepeira  grandiosa  alpina;  326,  Metepeira  celes- 
tun; 327,  Metepeira  vigilax;  328,  Metepeira  minima;  329,  Metepeira  uncata;  330,  Metepeira  pimungan;  331,  Metepeira  comanche. 
332-337.  Primarily  collected  during  the  northern  hemisphere  fall  season.  332,  Metepeira  lacandon;  333,  Metepeira  ctiilapae; 
334,  Metepeira  petatlan;  335,  Metepeira  spinipes;  336,  Metepeira  atascadero;  337,  Metepeira  triangularis. 


88         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No. 


tarsus  1.8,  tarsus  0.8.  Second  patella  and 
tibia  1.9  mm,  third  1.2,  fourth  1.7. 

Male  from  same  locality  as  female.  Male 
carapace,  dorsum,  venter,  sternum  darker 
and  more  contrasty  version  of  female 
(Figs.  296,  297).  All  legs  ringed.  Femur  I 
with  row  of  three  macrosetae  on  anterior 
side;  none  on  anteroventral  side.  Ratio  of 
eye  diameters:  posterior  medians  and  an- 
terior medians  0.9,  anterior  laterals  1.2, 
posterior  laterals  1.2.  Anterior  median 
eyes  separated  by  1.3  diameters,  posterior 
median  eyes  by  0.9,  anterior  median  eyes 
separated  from  anterior  laterals  by  1.2  di- 
ameters of  anterior  lateral  eyes,  lateral 
eyes  separated  by  2.3  their  diameters.  To- 
tal length  2.3  mm.  Carapace  1.2  mm  long, 
0.9  wide.  First  femur  1.4  mm,  patella  and 
tibia  1.3,  metatarsus  1.0,  tarsus  0.5.  Sec- 
ond patella  and  tibia  1.2  mm,  third  0.6, 
fourth  0.9. 

Diagnosis.  Unlike  other  members  of  the 
M.  minima  species  group,  M.  jamaicensis 
and  M.  pacifica  share  very  light  pigmen- 
tation and  the  embolus  in  both  species  is 
needle-shaped  (Figs.  286,  289,  291,  293, 
296,  298).  Unlike  M.  pacifica,  the  flagellae 
on  the  median  apophysis  of  M.  jamaicensis 
are  set  off  on  a  long,  thin  stalk  (Fig.  293) 
rather  than  a  short,  wide  one  (Fig.  286).  A 
ventral  view  of  the  epigynum  in  M.  jamai- 
censis shows  a  ridge  under  the  scape  that 
almost  forms  a  straight  line  (Fig.  295), 
compared  to  a  V-shape  (Fig.  288). 

Variation.  Average  body  length  of  elev- 
en females  examined  5  mm,  range  4.2  to 
6.1  mm.  Average  body  length  of  four 
males  examined  2.4  mm,  range  2.3  to  2.6 
mm. 

Natural  History.  Mature  specimens 
have  been  collected  in  July  through  March 
(Fig.  313). 

Distribution.  Primarily  in  Jamaica  and 
Haiti  (Map  15),  near  sea  level. 

Records    Examined.    BRITISH    WEST    INDIES 

Grand  Cayman  Island:  19°20'N,  81°10'W,  15.ii.l960 
(R.  A.  Lewin,  MCZ).  HAITI  Departement  de 
L'Ouest:  Port-au-Prince,  18°32'N,  72°20'W, 
19.vii.l955  (A.  F.  Archer,  AMNH),  20.vii.l955  (A.  F. 
Archer,  AMNH).  Dept.  de  L'Artibonite:  Saint-Marc, 


19°7'N,  72°42'W,  15,i.l913  (W.  M.  Mann,  MCZ).  JA- 
MAICA Cornwall:  Montego  Bay,  18°28'N,  77°55'W, 
l.iii.l984  (L.  E.  Schulten  Jr,  MCZ).  Middlesex:  3  mi 
E  Old  Harbor,  17°56'N,  77°10'W,  21.X.1957  (A.  M. 
Chickering,  MCZ);  Christiana,  18°10'N,  77°29'W, 
13.xi.l957  (A.  M.  Chickering,  MCZ),  15.vii.l960  (C. 
&  P  Vaurie,  AMNH),  17.vii.l960  (C.  &  P  Vaurie, 
AMNH);  Strawberry  Fields  near  Robin's  Bay  and 
Green  Casde,  18°17'N,  76°48'W,  23.iii.1972  (H.  W., 
L.  &  F  Levi,  MCZ),  25.iii.1972  (H.  W.,  L.  &  F.  Levi, 
MCZ),  26.iii.1972  (H.  W.,  L.  &  F  Levi,  MCZ).  Saint 
Ann:  Roaring  River,  18°24'N,  77°9'W  [?],  8.ii.l946 
(B.  Heineman,  AMNH);  Saint  Ann's  Bay,  18°26'N, 
77°8"W,  20.xi.l959  (A.  M.  Nadler,  AMNH).  Saint 
Catherine:  E  Green  Harbour,  S  slope  of  Healdishire, 
17°53'N,  76°51"W,  12.viii.l958  (A.  F.  Archer, 
AMNH).  St.  Andrews:  Ferry,  9/10  mi  on  Spanishtown 
Road,  18°2'N,  76°53"W,  26.vii.1955  (A.  F  Archer, 
AMNH).  Surrey:  Kingston,  Mona  Road,  pasture, 
17°59'N,  76°24"W,  lO.x.1957  (A.  M.  Chickering, 
MCZ);  Roselle  Falls,  24  mi  E  Kingston,  17°59'N, 
76°24"W,  29.X.1957  (A.  M.  Chickering,  MCZ);  Saint 
Andrew,  18°4'N,  76°45'W,  7.X.1957  (A.  M.  Chicker- 
ing, MCZ).  Trelawny:  Falmouth,  18°30'N,  77°39"W, 
20.vii.l960  (C.  &  P.  Vaurie,  AMNH).  Westinorland: 
Negril,  18°16'N,  78°21'W,  24.iii.1955  (A.  M.  Nadler, 
AMNH);  'Whitehouse,  18°4'N,  77°58"W,  26.iii.1955 
(A.  M.  Nadler,  AMNH). 

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Spiller,  D.  a.,  and  T.  W  Schoener.  1989.  Effect 
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Taczanowski,  L.  1878.  Les  araneides  du  Perou  Cen- 
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TULLGREN,  A.  1901.  Contribution  to  the  spider 
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Uetz,  G.  W,  and  C.  S.  Hieber.  1994.  Group  size 
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Uetz,  G.  W.,  C.  S.  Hieber,  E.  M.  Jakob,  R.  S.  Wil- 
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Uetz,  G.  W.,  and  M.  A.  Hodge.  1990.  Influence  of 
habitat  and  prey  availability  on  spatial  organiza- 
tion and  behavior  of  colonial  web-building  spi- 
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Uetz,  G.  W,  T.  C.  Kane,  and  G.  E.  Stratton.  1982. 
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Uetz,  G.  W,  T.  G.  Kane,  G.  E.  Stratton,  and  M. 
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ViERA,  C.  1986.  Comportamiento  de  captura  de  Me- 
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.  1989.  Caracteristicas  de  la  tela  orbicular  de 

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1992.  Gomparacion  de  telas  de  hembras  y 


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Wise,  D.  H.  1983.  Competitive  mechanisms  in  a 
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gia,  58:  1-9. 


INDEX 

Valid  taxon  names  are  printed  in  italics.  Page  numbers  in  bold  refer  to  illustrations;  those  in  italics  refer  to 
species  descriptions. 


acostai,  Metepeira  63 

Aculepeira  8 

alpina,  Metepeira  23,  24 

Amazonepeira  8 

Arachnidornyia  10 

Araneus  8 

arizonica,  Metepeira  9,  19,  65,  66 

atascadero,  Metepeira  9,  16,  18,  67,  69 

bani,  Metepeira  63 

biogeography  9,  10 

cajabamba,  Metepeira  9,  12,  17,  26,  31 

calamuchita,  Metepeira  14,  18,  39,  42 

cancriforrnis,  Gasteracantha  10 

celestun,  Metepeira  9,  10,  16,  19,  74,  75 

cereicola,  Metepeira  43 

chilapae,  Metepeira  9,  16,  19,  78,  79 

chilapica,  Metepeira  78 

Comanche,  Metepeira  14,  19,  60,  61 

compsa  group,  Metepeira  47 

compsa,  Aranea  48 

compsa,  Metepeira  7,  9,  10,  11,  14,  17,  48,  49 

crassipes,  Metepeira  7,  9,  10,  16,  19,  77,  79 

Cyrtophora  8,  9 

dakota,  Metepeira  23,  24 

daytona,  Metepeira  6,  7,  8,  9,  10,  12,  17,  20 

desenderi,  Metepeira  6,  10,  12,  17,  19,  21,  25 

digital  photography  4 

dominicana,  Metepeira  30 

dorsal  folium  6 

douglasi,  Metepeira  34 

ensenada,  Metepeira  71 


epigynum  8 

foxi  group,  Metepeira  19 
foxi,  Metepeira  8,  9,  10,  12,  17 
galatheae,  Araneus  43 
galatheae,  Epeira  43 
galatheae,  Metepeira  9,  14,  18,  43,  45 
glomerabilis,  Araneus  28 
glomerabilis,  Epeira  28 
glomerabilis,  Metazygia  28 
glomerabilis,  Metepeira  9,  12,  17,  28,  31 
gosoga,  Metepeira  14,  18,  59,  61 
grandiosa  alpina,  Metepeira  9,  12,  17,  24,  25 
grandiosa  grandiosa,  Metepeira  9,  10,  12,  17,  23,  25 
grandiosa  palustris,  Metepeira  9,  12,  17 
grandiosa,  Metepeira  23 
gressa,  Epeira  54 
gressa,  Metazygia  54 
gressa,  Metepeira  9,  18,  54,  57 
gressus,  Araneus  54 
grinnelli,  Metepeira  34 
habitats  9 

inca,  Metepeira  6,  14,  17,  18,  57,  58 
incrassata  group,  Metepeira  55 
incrassata,  Metepeira  8,  9,  10,  11,  16,  19,  68,  69 
inerma,  Metepeira  20 
jamaicensis,  Metepeira  9,  17,  19,  83,  86 
josepha,  Metepeira  77 
Kaira  6,  8 
karkii,  Araneus  46 
karkii,  Metepeira  9,  13,  18,  45,  46 
koepckeonim,  Araneus  6 


92         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  1 


labyrinthea  grinnelli,  Aranea  34 

labyrinthea  grinnelli,  Epeira  34 

labyiinthea  grinnelli,  Metepeira  34 

labyrintliea  group,  Metepeira  33 

labyrinthea,  Epeira  21 

labtjrinthea,  Metepeira  7,  9,  10,  14,  17,  32,  34,  46,  48 

labyrintheus,  Araneus  48 

lacandon,  Metepeira  9,  14,  17,  35,  37 

latigyna,  Metepeira  48 

lindae,  Arachnidoijia  10 

maija,  Metepeira  9,  14,  18,  56,  57 

measurements  5 

Mecynogea  8,  9 

median  apophysis  8 

Metepeira  5 

minima  group,  Metepeira  80 

minima,  Metepeira  9,  10,  12,  16,  19,  81,  83,  86 

nigriventris  group,  Metepeira  38 

nigriventris,  Araneus  38 

nigriventris,  Epeira  38 

nigriventris,  Metepeira  9,  10,  11,  12,  18,  38,  39 

ocosingo,  Mecynogea  10 

olmec,  Metepeira  9,  16,  19,  5.9,  61 

pacifica,  Metepeira  16,  19,  83,  84 

palomara,  Metepeira  23 

palp  8 

palustris,  Metepeira  23 

perezi,  Metepeira  48 

petatlan,  Metepeira  16,  19,  79,  80 

pimungan,  Metepeira  8,  9,  16,  19,  62,  65 

predation  10 

rayorae,  Arachnidomyia  10 

rectangula,  Epeira  32 


rectangula,  Metepeira  5,  6,  9,  12,  17,  32,  35 

rectangulata,  Metepeira  32 

revillagigedo,  Metepeira  16,  19,  73,  75 

roraima,  Metepeira  9,  12,  18,  49,  53 

Salei,  Epeira  68 

salei,  Metepeira  70 

sallei,  Aranea  70 

sallei,  Araneus  70 

santa,  Aranea  28 

scitulus,  Araneus  54 

seditiosa,  Epeira  54 

seditiosa,  Eustala  54 

seditiosa,  Metepeira  54 

seditiosus,  Araneus  54 

Singa  6 

species  groups  10 

spinipes,  Araneus  34 

spinipes,  Metepeira  5,  6,  8,  9,  14,  17,  34,  35 

suspended  retreat  8 

tarapaca,  Metepeira  7,  9,  14,  18,  39,  40 

triangularis,  Metepeira  9,  15,  18,  63,  65 

Trypargilum  10 

uncata,  Metepeira  9,  16,  19,  75,  76 

uncatus,  Araneus  76 

vaurieomm,  Metepeira  48 

Ventura  group,  Metepeira  71 

Ventura,  Metepeira  10,  16,  19,  69,  71 

vigilax  group,  Metepeira  26 

vigilax,  Araneus  30 

vigilax,  Epeira  29 

vigilax,  Metepeira  9,  10,  12,  17,  26,  30,  31 

virginensis,  Metepeira  48 

web  8 

Zygiella  6 


OF     THE 


seum 


(US  ISSN  0027-4100) 


Type  Specimens  of  Recent  Mammals 
in  the  Museum  of  Comparative  Zoology 


K.  M.  HELGEN  ANDT.  L.  McFADDEN 


1V1CZ 
LIBRARY 

JUL  0  3  2001 

HA^VARO 
y  N  L V  ii:^  R  -^  ^"^  ' 


HARVARD  UNIVERSITY 

CAMBRIDGE,  MASSACHUSETTS,  U.S.A. 


VOLUME  157,  NUMBER  2 
8  JUNE  2001 


(US  ISSN  0027-4100) 


PUBLICATIONS  ISSUED 

OR  DISTRIBUTED  BY  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 

HARVARD  UNIVERSITY 


Breviora  1952- 

bulletin  1863- 

Memoirs  1865-1938 

JoHNSONiA,  Department  of  Mollusks,  1941-1974 

Occasional  Papers  on  Mollusks,  1945- 

SPECIAL  PUBLICATIONS. 

1.  Whittington,  H.  B.,  and  W.  D.  I.  Rolfe  (eds.),  1963  Phylogeny  and 
Evolution  of  Crustacea.  192  pp. 

2.  Turner,  R.  D.,  1966.  A  Survey  and  illustrated  Catalogue  of  the  Tere- 
dinidea  (Mollusca:  Bivalvia).  265  pp. 

3.  Sprinkle,  J-.,  1973.  Morphology  and  Evolution  of  Blastozoan  Echino- 
derms.  284  pp. 

4.  Eaton,  R.  J.,  1974.  A  Flora  of  Concord  from  Thoreau's  Time  to  the 
Present  Day.  236  pp. 

5.  Rhodin,  A.  G.  J.,  and  K.  Miyata  (eds.),  1983.  Advances  in  Herpetology 
and  Evolutionary  Biology:  Essays  in  Honor  of  Ernest  E.  Williams. 

725  pp. 

6.  Angelo,  R.,  1990.  Concord  Area  Trees  and  Shrubs.  118  pp. 

Other  Publications. 

Bigelow,  H.  B.,  and  W.  C.  Schroeder,  1953.  Fishes  of  the  Gulf  of  Maine. 
Reprinted  1964. 

Brues,  C.T.,  A.  L.  Melander,  and  F.  M.  Carpenter,  1954.  Classification  of 
Insects.  {Bulletin  of  the  M.  C.  Z,  Vol.  108.)  Reprinted  1971. 

Creighton,  W.  S.,  1950.  The  Ants  of  North  America.  Reprinted  1966. 

Lyman,  C.  P.,  and  A.  R.  Dawe  (eds.),  1960.  Proceedings  of  the  First  In- 
ternational Symposium  on  Natural  Mammalian  Hibernation.  {Bulletin 
of  the  M.  C.  Z,  Vol  124.) 

Orinthological  Gazetteers  of  the  Neotropics  (1975-). 

Peter's  Check-list  of  Birds  of  the  World,  vols.  1-16. 

Proceedings  of  the  New  England  Zoological  Club  1899-1947.  (Complete 
sets  only.) 

Proceedings  of  the  Boston  Society  of  Natural  History. 

Price  list  and  catalog  of  MCZ  publications  may  be  obtained  from  Publica- 
tions Office,  Museum  of  Comparative  Zoology,  Harvard  University,  Cambridge, 
Massachusetts  02138,  U.S.A. 

This  publication  ha':  been  printed  on  acid-free  permanent  paper  stock. 

©The  President  and  Fellows  of  Harvard  College  2001. 


TYPE  SPECIMENS  OF  RECENT  MAMMALS  IN  THE  MUSEUM  OF 
COMPARATIVE  ZOOLOGY 

K.  M.  HELGEN  AND  T.  L  McFADDEN^ 

CONTENTS 


Abstract  -  94 

Introduction  94 

Authors  of  Type  Descriptions  94 

Definitions  and  Organization  96 

Taxa  Included  in  This  Catalogue  .._ 97 

Abbreviations  98 

Acknowledgments  98 

Paralectotype  Series 98 

Accounts  of  Type  Specimens  99 

Order  Didelphimoi"phia  99 

Family  Caluromyidae  99 

Family  Marmosidae  99 

Family  Didelphidae  100 

Order  Dasyuromoqohia  100 

Family  Dasyuridae  100 

Order  Peramelia  100 

Family  Peroiyctidae 100 

Order  Cingulata  101 

Family  Dasypodidae  101 

Order  Afrosoricida  101 

Family  Chiysochloridae  101 

Family  Tenrecidae  101 

Order  Rodentia  103 

Family  Aplodontidae  103 

Family  Sciuridae  103 

Family  Castoridae  109 

Family  Geomyidae  110 

Family  Heteromyidae 111 

Family  Dipodidae  112 

Family  Muridae  112 

Subfamily  AiAdcolinae  112 

Subfamily  Cricetinae  117 

Subfamily  Cricetomyinae  117 

Subfamily  Dendromurinae 117 

Subfamily  Gerbillinae  117 

Subfamily  Murinae  117 

Subfamily  Nesomyinae  120 

Subfamily  Otomyinae  120 

Subfamily  Sigmodontinae  121 

Family  Pedetidae  128 


^  Mammal  Department,  Museum  of  Comparative 
Zoology,  Harvard  University,  Cambridge,  Massachu- 
setts 02138. 


Family  Myoxidae  129 

Family  Bathyergidae  129 

Family  Erithizontidae 130 

Family  Dasyproctidae 130 

Family  Agoutidae  130 

Family  Octodontidae  130 

Family  Echimyidae  131 

Family  Capromyidae  131 

Order  Lagomoqiha  133 

Family  Ochotonidae  133 

Family  Leporidae  133 

Order  Scandentia  134 

Family  Tupaiidae 134 

Order  Primates  135 

Family  Indridae  135 

Family  Daubentoniidae  135 

Family  Galagonidae 135 

Family  Cebidae  135 

Family  Hylobatidae   136 

Family  Hominidae  136 

Order  Lipotyphla 136 

Family  Nesophontidae  136 

Family  Solenodontidae 137 

Family  Soricidae  137 

Family  Talpidae  140 

Order  Chiroptera 140 

Family  Pteropodidae  140 

Family  Emballonuridae  142 

Family  Nycteridae 142 

Family  Rhinolophidae  142 

Family  Mormoopidae  143 

Family  Phyllostomidae  143 

Family  Molossidae  145 

Family  Vespertilionidae 145 

Family  Thyi^opteridae  147 

Order  Artiodactyla 147 

Family  Tayassuidae  147 

Family  Monodontidae  148 

Family  Phocoenidae  148 

Family  Cervidae  148 

Family  Bovidae 149 

Order  Carnivora  149 

Family  Canidae  149 

Family  Ursidae  151 


Bull.  Mus.  Comp.  ZooL,  157(2):  93-181,  June,  2001         93 


94         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Family  Procyonidae  151 

Family  Mustelidae  152 

Family  Mephitidae 154 

Family  Viverridae  155 

Family  Heipestidae  155 

Family  Felidae  155 

Order  Cimolesta  156 

Family  Manidae 156 

References  157 

Index  171 

Abstract.  The  Mammal  Department  at  the  Mu- 
seum of  Comparative  Zoology  houses  name-bearing 
types  of  342  species-group  taxa  of  Recent  inammals. 
The  type  collection  consists  of  327  holotypes,  2  lec- 
totypes,  4  complete  syntype  series,  and  9  partial  syn- 
type  series.  This  catalogue  notes  information  on  the 
type  locality,  collector,  date  of  collection,  present  con- 
dition, original  publication,  and  synonyms  for  all 
name-bearing  types  in  the  Mammal  Department. 
Comments  on  the  taxonomic  and  historical  impor- 
tance of  many  type  specimens  are  included.  Lecto- 
types  for  a  number  of  taxa  are  designated  for  pur- 
poses of  taxonomic  consistency. 

INTRODUCTION 

"Some  of  them  are  more  or  less  historic 
specimens,"  wrote  the  great  mainiTialogist 
Glover  M.  Allen  in  1931,  "whose  location 
has  undoubtedly  in  many  cases  been  lost 
sight  of,  so  that  it  may  be  of  value  to  make 
the  present  record"  (1931:  230).  Allen, 
then  Curator  of  Mammals  at  the  Museum 
of  Comparative  Zoology  at  Harvard  Uni- 
versity, was  referring  to  the  type  speci- 
mens of  inammals  in  the  museum,  the 
subject  of  a  listing  he  published  that  year. 
Allen's  words  resound  with  even  greater 
truth  today  than  when  they  were  written 
70  years  ago.  Since  that  time,  the  inuse- 
um  s  holdings  of  mainmal  type  specimens 
have  grown  considerably.  Furthermore, 
type  specimens  of  ixiamiTials  housed  in  the 
MCZ  have  sometimes  been  credited  to  the 
collections  of  other  institutions  or  consid- 
ered to  no  longer  exist,  making  the  need 
for  a  new  catalogue  obvious. 

Type  specimens  are  biological  reference 
points  that  lend  objectivity  to  taxonomy 
and  are  thus  of  critical  importance  in  sys- 
teinatic  investigations;  accordingly,  "eveiy 
institution  in  which  name-bearing  types 
are  deposited  should  publish  lists  of  name- 
bearing  types  in  its  possession  or  custody" 


(International  Commission  on  Zoological 
NoiTienclature,  1999:  79).  Our  duty  in  this 
regard  is  thus  long  overdue,  and  this  cat- 
alogue should  sei've  to  fulfill  that  obliga- 
tion. Without  doubt,  it  will  alleviate  con- 
fusion in  the  scientific  coiximunity  as  to  the 
w^hereabouts  of  certain  specimens,  thought 
to  be  lost,  especially  those  type  speciinens 
that  were  acquired  from  Guillaume  Gran- 
didier.  In  addition,  it  should  bring  to  light 
information,  fonnerly  unavailable,  on  this 
museum  s  veiy  notable  holdings  of  name- 
bearing  types. 

All  taxonoinic  judgments  in  this  work, 
including  new  name  combinations  and  lec- 
totype  designations,  reflect  the  decision  of 
the  first  author  (KMH)  alone  and  should 
be  cited  accordingly. 

AUTHORS  OF  TYPE  DESCRIPTIONS 

Four  maminalogists  authored  the  over- 
whelming majority  of  names  based  on  type 
specimens  in  the  MCZ:  Glover  Allen,  Out- 
ram  Bangs,  Guillaume  Grandidier,  and 
Barbara  Lawrence.  Following  are  brief  bi- 
ographies of  these  four  outstanding  mam- 
malogists. 

Glover  M.  Allen  (1879-1942)  | 

Curator  of  Mammals  at  the  MCZ  from 
1924  until  his  death  in  1942,  Glover  Allen 
began  his  work  in  the  Mammal  Depart- 
ment in  1907.  He  was  known  as  a  careful, 
dependable  researcher  and  an  outstand- 
ing, patient  teacher.  His  goal  in  all  en- 
deavors was  to  increase  the  sum  total  of 
knowledge  about  the  mammals  of  the 
world.  Although  much  of  his  career  was 
spent  in  the  museum  studying,  as  he  put 
it,  "the  dried  remains  of  animals"  (Barbour 
et  al,  1943:  300),  he  also  traveled  widely 
throughout  the  world  collecting  specimens 
as  well  as  observing  and  learning  about  liv- 
ing animals.  He  once  commented  that  the 
actual  knowledge  of  living  creatures  could 
all  too  often  be  summed  up  by  saying 
"when  we  found  it,  it  ran  like  hell,  where- 
upon we  shot  it!"  (Barbour  et  al.,  1943: 
300).  He  wrote  prolifically;  his  bibliogra- 
phy of  publications  is  81  pages  long,  the 


Type  Specimens  of  Recent  Mammals  •  Helgen  and  McFadden        95 


first  of  which  he  pubHshed  when  he  was 
only  11  years  old  (Lawrence  1947a:  1). 
Holotypes  for  96  taxa,  which  Allen  de- 
scribed alone  or  with  colleagues,  are  de- 
posited in  the  MCZ. 

Outram  Bangs  (1863-1932) 

Outram  Bangs,  Curator  of  Mammals 
from  1899  to  1924,  "was  one  of  those  for- 
tunate mortals,  born  with  a  love  of  nature 
and  the  outdoors  which  rule  their  entire 
lives"  (Peters  1933:  265).  Bangs  authored 
135  MCZ  mammal  names  (only  one  co- 
authored!).  He  and  brother  Edward  began 
collecting  as  boys  using  slingshots  and 
horsehair  nooses.  He  was  an  early  ecolo- 
gist,  saving  Microtus  breweri  by  killing  the 
feral  cat  population  on  Massachusetts' 
Muskeget  Island  and  restocking  the  mouse 
from  a  tiny  islet  across  the  channel.  Al- 
though his  greatest  passion  was  the  natural 
history  of  birds,  he  was  also  fascinated  with 
mammals  and  served  as  curator  of  both 
departments  in  the  MCZ.  He  decided  to 
systematically  collect  the  mammals  of  east- 
ern North  America  in  about  1890.  He  be- 
gan by  trapping  in  New  England  and  later 
made  trips  to  the  southeastern  United 
States  and  Canada.  Other  collectors  assist- 
ed Bangs,  expanding  the  collection  area  to 
western  North  America  and  south  to  Cen- 
tral America.  His  precision  and  organiza- 
tion were  legendaiy,  and  today  his  mam- 
mal collection  of  more  than  10,000  speci- 
mens, donated  to  the  Museum  in  1899,  re- 
mains one  of  the  best  curated  and  most 
informative  in  the  department.  A  complete 
list  of  Bangs'  scientific  publications  was 
compiled  by  Porter  (1943). 

Guillaume  Grandidier  (1873-1957) 

French  explorer  and  scientist  Guillaume 
Grandidier,  son  of  naturalist  Alfred  Gran- 
didier, authored  and  coauthored  descrip- 
tions of  13  taxa  whose  types  were  donated 
to  the  MCZ  in  1947,  along  with  his  exten- 
sive personal  collection  of  Malagasy  mam- 
mals. The  collection  was  purchased  by 
Robert  Barbour  and  donated  to  the  MCZ 
in   honor   of  his    brother,    MCZ   director 


Thomas  Barbour.  Between  1898  and  1902, 
Grandidier  explored  the  center  and  south- 
ern portions  of  Madagascar,  collecting 
specimens  and  describing  the  geography 
of  the  area.  Through  his  writings  he 
brought  the  unique  fauna  of  this  remote 
region  to  the  attention  of  the  scientific 
world.  He  was  known  for  his  devotion  to 
the  careful  acquisition  of  knowledge, 
whether  it  was  geographical,  historical,  or 
scientific.  His  work  was  honored  by  both 
the  scientific  community  and  the  French 
government  (Chapus,  1953). 

Barbara  Lawrence  (1909-97) 

After  graduating  from  Vassar  College  in 
1931,  Barbara  Lawrence  became  a  volun- 
teer at  the  MCZ.  She  was  encouraged  by 
Dr.  Glover  Allen  to  do  her  own  research, 
and  in  the  late  1930s  she  made  field  trips 
to  the  Philippines  and  Sumatra  to  collect 
mammals.  In  1952  she  was  appointed  Cu- 
rator of  Mammals,  a  position  she  held  un- 
til her  retirement  in  1976.  Her  areas  of 
scientific  interest  were  many,  ranging  from 
echolocation  in  w^hales  to  zooarchaeology, 
as  well  as  the  more  traditional  mammalog- 
ical  pursuit  of  taxonomy.  "She  once  wrote 
'.  .  .  to  know  and  love  a  bit  of  the  world 
so  well  that  you  can  give  it  to  someone  else 
...  is  a  rare  talent'  "  (Rutzmoser,  1999: 
1049),  certainly  a  talent  Barbara  Lawrence 
had  in  abundance.  She  authored  or  coau- 
thored 20  MCZ  names. 

The  specimens  described  by  these  au- 
thors and  others  are  the  result  of  world- 
wide collecting  by  numerous  expeditions 
and  individuals,  listed  in  the  accounts  that 
follow.  A  handful  of  type  specimens  in  the 
MCZ  were  formerly  in  the  collection  of 
the  National  Museum  of  Natural  Histoiy 
in  Washington,  D.C.  According  to  Glover 
Allen,  these  were  obtained  via  exchanges 
"at  a  time  when  'duplicates'  were  more 
freely  disposed  of,  [and  would]  prove  to  be 
cotypes  or  even  actual  types"  (1931:  229). 
The  collections  of  the  Boston  Society  of 
Natural  Histoiy,  once  contained  in  the 
Boston  Museum  of  Science,  were  trans- 


96 


Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


ferred  to  the  Museum  of  Comparative  Zo- 
ology over  the  course  of  the  20th  century. 
Several  types  are  among  these  specimens, 
which  consist  primarily  of  mammals  from 
the  New  England  region. 

DEFINITIONS  AND  ORGANIZATION 
Type  Categories 

Type  specimens  are  categorized  as  one 
of  the  following: 

Holotype.  The  single  specimen  desig- 
nated as  name-bearer  in  the  original  pub- 
lished description  of  the  species-group  tax- 
on  (Article  73.1,  International  Commission 
on  Zoological  Nomenclature,  1999:  79). 

Syntype.  One  of  multiple  specimens  on 
which  a  species-group  name  is  equally 
based,  when  no  holotype  is  specified  in  the 
original  description  and  no  subsequent 
designation  of  a  lectotype  has  been  pub- 
lished (Article  73.2,  International  Com- 
mission on  Zoological  Nomenclature, 
1999:  81). 

Lectotype.  One  of  multiple  specimens 
upon  which  a  species-group  name  is  orig- 
inally based,  designated  in  a  publication 
subsequent  to  the  original  description  to 
become  the  unique  name-bearer  (Article 
74,  International  Commission  on  Zoologi- 
cal Nomenclature,  1999:  82). 

Neotype.  A  specimen  chosen  as  the 
name-bearer  of  a  species-group  taxon,  "if 
no  holotype,  lectotype,  syntype,  or  prior 
neotype  is  believed  to  exist"  (Article  75, 
International  Commission  on  Zoological 
Nomenclature,  1999:  84).  There  are  no 
neotypes  in  the  MCZ  mainmal  collection. 

Paratype.  A  specimen  other  than  the 
holotype  (if  designated  originally)  that  is 
mentioned  in  the  original  description  of  a 
species-group  taxon.  Paratypes  of  species- 
group  taxa  whose  holotypes  are  housed  in 
other  institutions  are  not  mentioned  in  this 
catalogue,  although  inany  such  specimens 
exist  in  this  museum. 

Paralectotype.  If  no  holotype  is  desig- 
nated, a  specimen  other  than  the  lectotype 
(if  designated  subsequently)  that  is  men- 
tioned in  the  original  description  of  a  spe- 


cies-group taxon.  Paralectotypes  of  spe- 
cies-group taxa  whose  lectotypes  are 
housed  in  other  institutions  are  discussed 
below,  before  the  accounts  for  name-bear- 
ing types. 

Locality 

This  categoiy  includes  country,  second- 
level  geopolitical  division  (state,  depart- 
ment, territory,  province,  or  district),  col- 
lection site,  and  altitude  where  available. 
The  geographic  name  given  for  a  type  lo- 
cality is  that  found  in  the  original  publi- 
cation. When  the  description  of  the  col- 
lection locality  does  not  include  current 
geopolitical  divisions,  that  information  is 
provided  in  parentheses.  If  an  original 
name  is  no  longer  used,  an  equals  sign  ( = ) 
is  included  within  the  parentheses  to  des- 
ignate an  equivalent  modern  name.  Where 
altitude  was  originally  given  in  feet,  it  has 
been  converted  to  meters  and  included  in 
parentheses. 

Sources  used  for  the  current  names  are 
from  the  most  current  available  Gazetteers 
of  the  United  States  Board  on  Geographic 
Names,  the  10th  comprehensive  edition  of 
the  Times  Atlas  of  the  World,  the  Colum- 
bia Gazetteer  of  the  World,  and  the  Or- 
nithological Gazetteers  of  South  America. 

Account  Organization 

The  type  locality,  collector,  date  of  col- 
lection, and  present  condition  of  each 
specimen  are  noted.  The  publication  of 
the  original  description  is  cited  for  each 
specimen.  Many  names  have  changed  in 
rank  or  synonymy  since  their  origin;  in 
these  cases,  the  name  by  which  a  taxon  is 
known  today  is  noted,  with  a  citation  of  the 
publication  in  which  that  name  combina- 
tion was  first  employed  for  that  taxon. 
Comments  are  offered  for  most  entries  to 
provide  additional  information  or  to  dispel 
potential  sources  of  confusion. 

The  format  of  this  catalogue  is  largely 
borrowed  from  the  most  recent  type  cat- 
alogue of  inammals  in  the  American  Mu- 
seum of  Natural  History  (Lawrence, 
1993).  For  systematic  consistency,  the  tax- 


Type  Specimens  of  Recent  Mammals  •  Helgen  and  McFadden        97 


onomic  judgments  at  the  species-level  by 
the  authors  of  the  chapters  in  Mammal 
Species  of  the  World  (Wilson  and  Reeder, 
1993)  are  largely  adhered  to.  However,  in 
many  cases  alternate  views  are  explored, 
and  subsequent  work  by  other  authors  is 
noted.  The  sequence  of  mammalian  orders 
presented  here  is  as  follows:  Didelphimor- 
phia,  Dasyuromorphia,  Peramelia,  Cingu- 
lata,  Afrosoricida,  Rodentia,  Lagomoi-pha, 
Scandentia,  Primates,  Lipotyphla,  Chirop- 
tera,  Artiodactyla,  Carnivora,  and  Cimoles- 
ta.  This  sequence  represents  the  ongoing 
understanding  of  higher  mainmalian  rela- 
tionships being  produced  by  research  in 
molecular  phylogenetics  (Waddell  et  al., 
1999)  as  well  as  paleontological  studies 
(McKenna  and  Bell,  1997).  A  number  of 
these  names  are  rather  nontraditional;  use 
of  the  names  Cingulata  (for  armadillos) 
and  Cimolesta  (for  pangolins)  at  ordinal 
rank  follows  McKenna  and  Bell  (1997).  Af- 
rosoricida is  used  for  an  order  including 
tenrecs  and  golden-moles  (following  Stan- 
hope et  al,  1998:  9971-9972).  We  use  the 
ordinal  name  Lipotyphla  in  a  restricted 
sense  to  refer  to  the  Recent  families  Ne- 
sophontidae,  Solenodontidae,  Soricidae, 
and  Talpidae  (others  have  used  the  term 
"Eulipotyphla,"  e.g.,  Waddell  et  al,  1999). 
Cetaceans  are  included  here  within  the  or- 
der Artiodactyla  (an  assemblage  often  re- 
ferred to  as  "Cetartiodactyla"  in  recent  lit- 
erature; see  Graur  et  al.,  1997). 

To  avoid  unnecessaiy  complexity,  or- 
ders, families,  and  genera  are  the  only 
ranks  above  the  level  of  species  that  are 
listed,  except  for  the  large  family  Muridae, 
for  which  subfamilial  distinctions  are  pro- 
vided, in  alphabetical  order.  Within  each 
order,  the  sequence  of  families  generally 
follows  Simpson  (1945),  but  Wilson  and 
Reeder's  (1993)  order  of  rodent  families 
and  Simmons'  (1998:  12)  arrangement  of 
the  bats  are  observed.  Within  each  genus, 
taxa  are  presented  in  alphabetical  order  by 
original  name. 

The  format  of  this  catalogue  is  as  fol- 
lows, with  all  the  following  information 
provided  when  possible: 


Original  binomen.  Name  of  describer,  date 

of  description. 

Citation  of  original  publication. 

= Presently  used  name,   if  different  from 

original.  Citation  of  publication  in  which 

this  name  combination  was  initially  used 

for  this  taxon. 

Type  Category.  Number  of  specimen.^ 
Preparation  of  specimen  (skin,  skull,  alco- 
hol, etc.),  age  and  sex. 

Locality.  Type  locality.  Date  of  collec- 
tion. 

Collector.  Name  of  collector.  Original 
number  of  specimen. 

Condition.  Current  condition  of  the 
type  material. 

Type  Series.  Any  paratypes,  paralecto- 
types,  or  additional  syntypes  in  existence 
are  mentioned,  with  their  preparation,  sex, 
and  age. 

Comments.  Additional  comments  re- 
garding the  systematic  status  or  the  histoiy 
of  the  specimen. 

TAXA  INCLUDED  IN  THIS  CATALOGUE 

Unlike  Allen's  original  type  catalogue,  fos- 
sil mammals  are  not  considered  here,  but 
type  specimens  of  Recent  mammals  known 
only  from  subfossil  remains  are  discussed. 

Several  type  specimens  of  Recent  mam- 
mals that  were  included  in  Allen's  cata- 
logue are  not  considered  here  as  name- 
bearing  types.  A  number  of  syntype  series 
in  the  MCZ  have  since  Allen's  time  been 
rendered  paralectotypes  by  the  designa- 
tion of  a  lectotype  preserved  in  another 
institution;  these  are  discussed  in  the  sec- 
tion below  on  paralectotypes.  Additionally, 
MCZ  14929,  listed  as  a  "cotype"  of  Nyc- 
teris  revoili  Robin,  1881  by  G.  M.  Allen 
(1931:  235),  is  a  paratype  rather  than  a 
syntype  and  is  not  considered  here. 


-  Mammal  specimens  in  the  MCZ  bear  any  of  three 
kinds  of  numbers.  A  number  preceded  by  "MCZ"  can 
be  found  in  the  general  collection  in  the  Mammal 
Department.  A  number  preceded  by  a  "B"  is  part  of 
the  collection  of  E.  A.  and  O.  Bangs,  also  housed  in 
the  Mammal  Department.  A  number  preceded  by 
"VP"  designates  that  the  specimen  is  stored  in  the 
Vertebrate  Paleontology  Department. 


98 


Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


One  other  enigmatic  specimen  in  the  col- 
lection deseives  some  mention.  When  it  was 
received  by  die  Mammal  Department,  Guil- 
laume  Grandidiers  personal  collection  con- 
tained specimens  marked  as  types  for  16 
taxa,  described  by  either  Grandidier  or  M. 
L.  Lavauden.  Fifteen  of  these  holotypes  are 
discussed  witliin  this  catalogue  in  separate 
accounts.  The  remaining  specimen,  a  fioiit 
bat,  MCZ  45073,  is  marked  exactly  like  the 
odier  type  specimens,  and  both  its  sldn  and 
its  skull  tags  bear  the  name  "Eidolon  saka- 
lava  nov.  spec,  G.  Grandidier"  in  die  script- 
ed handwriting  typical  of  Grandidiers  spec- 
imens. The  locality  given  for  the  specimen 
is  Ankavandra,  in  west-central  Madagascar. 
Though  this  specimen  has  been  curated  as 
a  type  specimen,  I  (KMH)  can  find  no  pub- 
lished description  of  diis  taxon  or  any  ref- 
erence to  diis  name  in  the  literature.  This 
name  is  dierefore  invalid,  barring  a  discov- 
eiy  in  the  future  diat  it  was  indeed  pub- 
lished and  has  since  been  overlooked.  What- 
ever die  published  status  of  the  name,  it  ap- 
pears to  me  diat  diis  specimen  should  be 
considered  as  a  young  specimen  of  Eidolon 
dupreanum  rather  dian  a  distinct  species, 
and  discussion  of  this  mysterious  binomial 
here  should  in  no  way  be  constiaied  as  a 
formal  description  of  a  new  taxon. 

ABBREVIATIONS 

Abbreviations  are  used  in  the  text  to 
designate  the  following  institutions. 

AMNH  American  Museum  of  Natural 
Histoiy,  New  York 

BMNH  Natural  Histoiy  Museum,  Lon- 
don 

BSNH  Boston  Society  of  Natural  His- 
tory, Boston 

FMNH  Field  Museum  of  Natural  His- 
tory, Chicago 

MNHN  Museum  National  d'Histoire 
Naturelle,  Paris 

MVZ  Museum  of  Vertebrate  Zoology, 

Berkeley,  California 

RMNH  Rijksmuseum  van  Natuurlijke 
Historic,  Leiden 


USNM 


YPM 


National   Museum   of  Natural 
Histoiy  Washington,  D.C. 
Yale  Peabody  Museum  of  Nat- 
ural History,  New  Haven,  Con- 
necticut 


ACKNOWLEDGMENTS 

We  would  like  to  thank  the  faculty  and 
staff  of  the  Mammal  Department  at  the 
MCZ — A.  W.  Crompton,  Andrew  Biewe- 
iier,  Judith  Chupasko,  Jane  Harrison,  and 
especially  Maria  Rutzmoser  for  access  to 
the  type  specimens.  Maiy  Sears,  Timothy 
McNeece,  and  Ronnie  Broadfoot  at  the 
Ernst  Mayr  Libraiy,  HaiA^ard  University, 
were  extremely  helpful  in  pinning  down 
difficult  references.  We  also  thank  Alison 
Pine  for  allowing  us  to  make  use  of  the 
MCZ  Ornithology  Department  locality 
references;  Carolyn  Kirdahy  of  the  Boston 
Museum  of  Science,  w^lio  granted  us  ac- 
cess to  the  archives  of  the  Boston  Society 
of  Natural  History;  and  the  many  individ- 
uals associated  with  other  museums  who 
assisted  by  answering  any  questions  that 
arose.  Finally,  we  are  thankful  for  the  sug- 
gestions we  received  from  two  anonymous 
reviewers,  which  helped  us  improve  this 
catalogue. 

PARALECTOTYPE  SERIES 

Hesperomys  eremicus  Baird,  1858  =Peromyscus  er- 
emicus  eremicus  (Baird,  1858).  Lectotype,  USNM 
2575,  designated  by  Osgood  (1909:  241).  MCZ 
4310  and  5273  are  paralectotypes. 

Neotoma  fuscipes  Baird,  1858  =Neotoinafuscipesfus- 
cipes  Baird,  1858.  Lectotype,  USNM  22026,  des- 
ignated by  Lyon  and  Osgood  (1909:  99).  MCZ  4336 
and  5264  are  paralectotypes. 

Mus  bairdii  Hoy  and  Kennicott,  1857  =Peromyscus 
maniculatus  bairdii  (Hoy  and  Kennicott,  1857). 
Lectotype,  number  750  in  the  Collection  of  the 
Academy  of  Natural  Sciences  of  Philadelphia,  des- 
ignated by  Osgood  (1909:  80).  MCZ  8073  is  a  par- 
alectotype. 

Pteropus  lanigera  [sic]  H.  Allen,  1890  =Pteropus  in- 
stdaris  Hombron  and  Jacquinot,  1842.  The  cor- 
rected spelling  of  the  original  name  is  Pteropus  lan- 
iger  (see  Andersen  1912:  297).  H.  Allen  based  the 
description  of  Pteropus  laniger  on  two  syntypes, 
USNM  19066  (skin)/37815  (skull)  and  MCZ  7023, 
a  skin.  Andersen  (1912:  297-298)  was  unaware  of 


Type  Specimens  of  Recent  Mammals  •  Helaen  and  McFadden 


99 


the  whereabouts  of  this  latter  specimen  and  based 
his  evaluation  of  the  systematic  status  of  P.  laniger 
solely  on  the  USNM  specimen  in  his  taxonomic 
review  of  the  Megachiroptera.  Because  it  is  rep- 
resented by  both  a  skin  and  a  skull  in  good  con- 
dition and  because  it  has  been  used  in  past  taxo- 
nomic treatments,  USNM  19066/37815  is  hereby 
designated  as  the  lectotype  of  Pteropus  laniger  to 
ensure  consistency  between  past  and  future  taxo- 
nomic treatments  of  this  name.  MCZ  7023  is  thus 
a  paralectotype.  The  type  locality  of  laniger  is  the 
Caroline  Islands,  as  emended  by  Andersen  (1912: 
298),  not  Samoa,  as  originally  described. 

Sciunis  castanotus  Baird,  1855  and  Sciurus  castan- 
onotus  Baird,  1858  =Scmnis  aherti  aherti  Wood- 
house,  1853.  Baird  (1858:  266)  noted  that  the  name 
castanotus,  used  in  his  original  description  of  this 
taxon,  was  a  misprint  for  castanonotus.  Lectotype, 
USNM  121/1107,  designated  by  Lyon  and  Osgood 
(1909:  183).  Though  not  explicitly  stated  by  Lyon 
and  Osgood,  this  specimen  should  serve  as  a  lec- 
totype for  both  names  (Sciunis  castanotus  Baird, 
1855  and  Sciunis  castanonotus  Baird,  1858),  for 
ttixonomic  consistency.  MCZ  4692  is  a  paralecto- 
type. 

Spennophiliis  obsoletus  Kennicott,  1863  =Spenno- 
philus  spilosoma  obsoletus  Kennicott,  1863.  Lec- 
totyjoe,  USNM  3222/27998,  designated  by  A.  H. 
Howell  (1938:  130).  See  G.  M,  Allen  (1931:  252) 
for  a  list  of  paralectotypes  (then  considered  synty- 
pes)  in  the  MCZ. 

Spennophiliis  parnji  van  kodiacensis  J.  A.  Allen,  1874 
=  Spennophiliis  parnji  kodiacensis  J.  A.  Allen, 
1874.  Lectotype,  USNM  9242/38543,  designated 
by  A.  H.  Howell  (1938:  103).  See  G.  M.  Allen 
(1931:  252)  for  a  list  of  paralectotypes  (then  con- 
sidered syntypes)  in  the  MCZ. 

Spennophiliis  tridecemlineatus  van  pallidus  ].  A.  Al- 
len, 1874  =  Spennophiliis  tridecemlineatus  pallidus 
J.  A.  Allen,  1874.  Lectotype,  USNM  16237,  des- 
ignated by  A.  H.  Howell  (1938:  112).  See  G.  M. 
Allen  (1931:  253)  for  a  Ust  of  paralectotypes  (then 
considered  syntypes)  in  the  MCZ. 

Tamias  quadrivittatus  var.  pallidus  J.  A.  Allen,  1874 
=  Tamias  minimus  pallidus  J.  A.  Allen,  1874.  Lec- 
totype, USNM  11656/38311,  designated  by  Gary 
(1906:  88).  G.  M.  Allen  (1931:  255)  provided  a  hst 
of  paralectotypes. 

ACCOUNTS  OF  NAME-BEARING  TYPE 
SPECIMENS 

Order  DIDELPHIMORPHIA  Gill,  1872 

Family  CALUROMYIDAE  Kirsch  and  Reig, 
1977 

Genus  CALUROMYSJ.  A.  Allen,  1900 

Philander  cicur  Bangs,  1 898k 


Proc.  Biol.  Soc.  Washington,  12:  161,  10 

August. 

=  Caluromys  lanatus  cicur  {Bangs,  1898). 

See  Cabrera  (1958:  2). 

Holotype.  B8114.  Skin  and  skull.  Adult  female. 

Locality.  Colombia:  (Magdalena),  Santa  Marta 
Mountains,  Pueblo  Viejo,  8,000  ft  (2,440  m).  27 
March  1898. 

Collector.  W.  W.  Brown,  Jr.  Original  number  123. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Tijpe  Series.  3  paratypes;  B8036,  skin  and  skull, 
adult  male;  B8115,  skin  and  skull,  adult  male; 
B8116,  skin  and  skull,  adult  male. 

Family  MARMOSIDAE  Hershkovitz,  1992 

Genus  MARMOSA  Gray,  1821 

Marmosa  robinsoni  Bangs,  18981 

Proc.  Biol.  Soc.  Washington,  12:  95,  30 

April. 

=  Marmosa  robinsoni  robinsoni  Bangs, 

1898.  See  Cabrera  (1958:  24). 

Holotype.  B7749.  Skin  and  skull.  Adult  male. 

Locality.  Venezuela:  (Nueva  Esparta),  Margarita 
Island.  12  July  1895. 

Collector.  W.  Robinson.  Original  number  506. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  2  paratypes,  both  in  the  USNM; 
USNM  63209,  skin  and  skull,  adult  male;  USNM 
63210,  skin  and  skull,  adult  female. 
Comments.  M.  robinsoni  was  considered  a  valid 
species  by  Gardner  (1993:  18)  and  Nowak  (1999: 
21). 

IVIarmosa  mitis  Bangs,  1 898k 

Proc.  Biol.  Soc.  Washington,  12:  162,  10 

August. 

=  Marmosa  robinsoni  robinsoni  Bangs, 

1898.  See  Cabrera  (1958:  24). 

Holotype.  B8123.  Skin  and  skull.  Adult  male. 

Locality.  Colombia:  (Magdalena),  Santa  Marta 
Mountains,  Pueblo  Viejo,  8,000  ft  (2,440  m).  25 
March  1898. 

Collector.  W.  W.  Brown,  Jr.  Original  number  91. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Tijpe  Series.  26  paratypes;  B8117-B8122,  B8124- 
B8143;  all  represented  by  skin  and  skull,  15  fe- 
males and  11  males.  5  paratypes  are  no  longer  in 
the  MCZ  (B8118  is  at  Wellesley  College,  B8124 
and  B8141  are  at  FMNH,  and  B8136  and  B8138 
are  at  USNM). 


100 


Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Marmosa  fulviventer  Bangs,  1901b 
Amer.  Nat.,  35:  632,  22  August. 
=  Marmosa  robinsoni  fulviventer  Bangs, 
1901.  See  Handley  (1966:  775). 

Holotype.  B8435.  Sldn  and  skull.  Adult  male. 

Locality.  Panama:  (Panama),  Gulf  of  Panama,  San 
Miguel  Island.  28  April  1900. 

Collector.  W.  W.  Brown,  Jr.  Original  number  123. 
Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  3  paratypes;  B8436,  skin  and  skull,  fe- 
male (exchanged  to  FMNH  in  1931);  B8437,  skin 
and  skull,  female;  B8438,  skin  and  skull,  male. 
Comments.  M.  r.  fulviventer  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  14)  and  O'Connell  (1983: 
1). 

Family  DIDELPHIDAE  Gray,  1821 

Genus  D/DELPH/S  Linnaeus,  1758 

Didelphis  marsupialis  particeps  Goldman, 

1917 

Proc.  Biol.  Soc.  Washington,  30:  107,  23 

May. 

Holotype.  B8439.  Skin  and  skull.  Adult  male. 

Locality.  Panama:  (Panama),  Gulf  of  Panama,  San 

Miguel  Island.  8  May  1900. 

Collector.  W.  W.  Brown,  Jr.  Original  number  165. 

Condition.  Skin  and  skull  complete. 

Type   Series.    1   paratype;   B8440,   skin   and  skull, 

adult  female. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 

(1981:  4). 

Didelphis  virginiana  pigra  Bangs,  1 898b 
Proc.  Boston  Soc.  Nat.  Hist.,  28:  172,  15 
March. 

Holotype.  B3500.  Sldn  and  skull.  Adult  female. 
Locality.  (United  States):  Florida,  Brevard  Gounty, 
Oak  Lodge,  East  Peninsula  opposite  Micco.  31  Jan- 
uaiy  1895. 
Collector.  O.  Bangs. 
Condition.  Skin  and  skull  complete. 
Type  Series.   11  paratypes;  all  represented  by  skin 
and  skull,  most  still  in  the  MGZ. 
Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  5)  and  McManus  (1974:   1).  The  type  de- 
scription lists  31  January  1896  as  the  date  of  col- 
lection,  but  the   date  is  written   as   "January  31, 
1895"  on  the  original  specimen  label  and  in  Bangs" 
accession  catalogue. 

Order  DASYUROIVIORPHIA  Gill,  1872 
Family  DASYURIDAE  Goldfuss,  1820 
Genus  ANTECHINUS  Macleay,  1841 


Antecfiinus  may  eh  misim  Tate,  1947 

Bull.  Amer.  Mus.  Nat.  Hist.,  88:  130,  20 

February. 

= Antecfiinus  naso  misim  Tate,  1947.  See 

Laurie  and  Hill  (1954:  7). 

Holotype  MGZ  29924.  Skin  and  skull.  Adult  male. 
Locality.   Papua  New  Guinea:   Morobe  Province, 
Mount  Misim  (  =  Missim),  5,850  ft  (1,784  m).  14 
April  1933. 
Collector  H.  Stevens. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  1  paratype;  MGZ  29923,  sldn  and 
skull,  adult  female. 

Comments.  Tate,  in  the  original  description,  erro- 
neously listed  24  April  1933  as  the  date  of  collec- 
tion. A.  n.  misim  was  retained  as  a  valid  subspecies 
by  Flanneiy  (1995a:  80),  who  also  noted  that  the 
New  Guinean  species  assigned  to  the  genus  Ante- 
chinus  are  not  closely  related  to  the  Australian  spe- 
cies of  that  genus  and  will  be  reassigned  at  the 
generic  level  pending  a  full  taxonomic  revision  of 
the  group. 

Genus  IVIYOICTIS  Gray,  1858 

Myoictis  melas  wavicus  Tate,  1 947 

Bull.  Amer.  Mus.  Nat.  Hist.,  88:  140,  20 

February. 

=  Myoictis  melas  wallacei  Gray,  1858.  See 

Flanneiy  (1990:  56). 

Holotijpe.  MGZ  28082.  Skin  and  skull.  Adult  male. 
Locality.   Papua  New  Guinea  (S.  Morobe),  Wau, 
3,800  ft  (1,159  m).  27  March  1932. 
Collector  H.  Stevens.  Original  number  1. 
Condition.  Sldn  and  skull  complete. 
Type  Series.  Holot}'pe  only. 

Comments.  Stevens'  original  field  label  bears  the 
following  lament — "To  my  intense  anguish  I  failed 
to  retrieve  the  female,  shot  on  recumbent,  decayed 
log  in  undergrowth." 

Order  PERAMELIA  Ameghino,  1889 

Family  PERORYCTIDAE  Groves  and 
Flannery,  1990 

Genus  ECHYMIPERA  Lesson,  1842 

Suillomeles  hispida  G.  M.  Allen  and 

Barbour,  1909 

Proc.  New  England  Zool.  Club,  4:  44,  12 

July. 

=  Echymipera  kalubu  kalubu  (Fischer, 

1829).  See  Laurie  and  Hill  (1954:  11). 

Holotype.  MGZ  7006.  Skin  and  skull.  Adult. 


Type  Specimens  of  Recent  Mammals  •  Helaen  and  McFadden 


101 


Localitij.  (Indonesia):  Dutch  New  Guinea  (  =  Ii-ian 

Jaya),  Doreh  Bay,  Manokwari,  "not  far  from  the 

foot  of  Mt.  Arfak."  23  Februaiy  1907. 

Collector.  T.  Barbour. 

Condition.   Skin  complete.  Skull  partial  (occipital 

region  and  two  posterior  upper  molars  on  each  side 

missing). 

Type  Serie.s.  Holotype  only. 

Comments.  S.  hispida  is  the  type  species  of  Suil- 

lomeles  G.  M.  Allen  and  Barbour,  1909. 


Echymipera  rufescens  austral  is  Tate,  1948 
Bull.  Amer.  Mus.  Nat.  Hist.,  92:  334,  25 
November. 


Holotype.  MCZ  29214.  Skin  and  skull.  Adult  male. 
Locality.   Australia:  Queensland,  Cape  York,  near 
Coen,  east  slope  of  Mcllwraith  Ranges.  Rocky  Riv- 
er, "Rocky  Scrub."  20  June  1932. 
Collector.  R  J.  Darlington,  Jr.,  Hanard  Australian 
Expedition.  Original  number  209. 
Condition.    Skin   complete,   but  tip  of  tail  worn. 
Skull  complete. 
Type  Series.  Holotype  only. 

Comments.  Tate  (1952:  582)  reported  that  the  ex- 
act type  locality  is  "in  the  dense  rain  forests  of  the 
upper  Nesbit  River  on  the  east  slopes  of  the 
Mcllwraith  Range."  Retained  as  a  valid  subspecies 
by  Flanneiy  (1995a:  111).  Darlington  noted  in  his 
field  notebook,  regarding  this  specimen,  "Animal 
about  the  fattest  I  have  skinned — I  shall  dream  of 
it!" 


Order  CINGULATA  llliger,  1911 

Family  DASYPODIDAE  Gray,  1821 

Genus  D/ASVPL/S  Linnaeus,  1758 

Dasypus  novemcinctus  hoplites  G.  M. 

Allen,  1911a 

Bull.  Mus.  Comp.  Zool.,  54:  195,  July. 


Holotype.    MCZ   8116.    Skin,    skull,   and  postcranial 
skeleton.  Adult  female. 

Locality.  Grenada:  hills  back  of  Gouyave.  7  Sep- 
tember 1910. 

Collector  G.  M.  Allen.  Original  number  26. 
Condition.    Skin,    skull,    and   postcranial    skeleton 
complete. 

Type  Series.  2  paratypes;  MCZ  8117,  skin  and  skull, 
adult  male;  MCZ  8118,  skin  and  skull,  adult  male. 
Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  283)  and  McBee  and  Baker  (1982:  1). 


Order  AFROSORICIDA  Stanhope  et  a!., 
1998 

Family  CHRYSOCHLORIDAE  Gray,  1825 

Genus  CHRYSOCHLORIS  Lacepede, 
1799 

Chlorotalpa  tropicalis  G.  M.  Allen  and 

Loveridge,  1927 

Proc.  Boston  Soc.  Nat.  Hist.,  38:  418,  23 

December. 

=  Chrysochloris  stuhlmanni  tropicalis  (G. 

M.  Allen  and  Loveridge,  1927).  See 

Meester  (1974:  3). 

Holotype.  MCZ  22435.  Skin  and  skull.  Adult  female. 
Locality.   Tanganyika  Territory  (=Tanzania):  Ulu- 
gui"u  Mountains,  Bagilo.  5  October  1926. 
Collector  A.  Loveridge. 

Condition.   Skin  complete.  Skull  slightly  damaged 
(coronoid  processes  missing  from  both  mandibular 
rami).  Mandible  disarticulated. 
Type  Series.  Holotyj^De  only. 

Comments.  Included  in  Chrysochloris  stuhlmanni 
by  Meester  (1974:  3)  but  recognized  as  distinct  by 
Simonetta  (1968:  42).  Hutterer  (1993:  75)  noted 
that  the  systematic  status  of  tropicalis  merits  fur- 
ther study. 

Family  TENRECIDAE  Gray,  1821 

Genus  GEOGALE  Milne-Edwards  and  A. 
Grandidier,  1872 

Cryptogale  australis  G.  Grandidier,  1928 

Bull.  Mus.  Hist.  Nat.  Paris,  34:  64,  26 

Januaiy. 

=  Geogale  aurita  Milne-Edwards  and  A. 

Grandidier,  1872.  See  Genest  and  Petter 

(1975:  3). 

Holotype.  MCZ  45057.  Skull  fragments. 

Locality.  Madagascar:  (Toliary),  south  of  Fort  Dau- 
phin (=Tolanaro),  Andrahomana  grotto.  1927. 
Collector  R.  Decary. 

Condition.   MCZ  45047  includes  18  partial  crania 
and  8  mandibular  rami.  Fragmentary. 
Type  Series.  All  the  type  material  of  C  australis 
bears  a  single  accession  number. 
Comments.  C.  australis  is  the  type  species  of  the 
genus  Cn/pto^ale  G.  Grandidier,  1928. 

Geogale  aurita  orientalis  G.  Grandidier 

and  Petit,  1930 

Faune  des  Colonies  Frangaises,  4:  446. 

Holotype.  MCZ  45660.  Body  in  alcohol,  cranium  sep- 
arate. 


102         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Locality.  Madagascar:  (Toamasina),  east  coast,  Fe- 
iierive  (  =  Fenoarivo  Atsinanana).  April  1928. 
Collector.  R.  Decaiy.  Original  number  12. 
Condition.  Alcoholic,  cranium  complete. 
Type  Series.  Holotype  only. 

Comments.  Retained  as  a  valid  subspecies  by  Ge- 
nest  and  Fetter  (1975:  3).  The  holotype  of  G.  a. 
orientalis  seems  to  be  the  only  record  of  Geogale 
aurita  from  the  east  coast  of  Madagascar. 

Genus  /W/CROG/\/.E  Thomas,  1882 

Microgale  decaryi  G.  Grandidier,  1928 

Bull.  Mus.  Hist.  Nat.  Paris,  34:  69,  26 

January. 

=  Microgale  principula  Thomas,  1926.  See 

MacPhee  (1987a:  9). 

Holotype.  MCZ  45049.  Cranium.  Adult. 

Locality.  Madagascar:  (Toliary),  south  of  Fort  Dau- 
phin (=Tolanaro),  Andrahomana  caves.  1926. 
Collector.  R.  Decary. 

Condition.  Cranium  partial  (back  of  cranium  miss- 
ing posterior  to  parietals). 

Type  Series.  Paratype  material  consists  of  MCZ 
45408,  which  includes  3  partial  crania  and  5  man- 
dibular rami,  and  MAD- 1649,  a  partial  skull  in  the 
collections  of  the  Institut  de  Paleontologie, 
MNHN. 

Microgale  drouhardi  G.  Grandidier,  1934 
Bull.  Mus.  Hist.  Nat.  Paris,  6:  474,  29 
November. 

Holotype.  MCZ  45034.  Body  in  alcohol,  skull  extract- 
ed. Juvenile  female. 

Locality.  Madagascar:  (Antsiranana),  east  coast,  Di- 
ego-Suarez  (=  Antsiranana).  May  1934. 
Collector  M.  E.  Drouhard.  Original  number  A. 
Condition.  Alcoholic,  skull  complete. 
Type  Series.  6  paratypes;  MCZ  46007-MCZ  46012, 
all  in  alcohol.  MCZ  46017,  represented  by  a  skull 
and  postcranial  skeleton,  was  collected  at  the  same 
time  and  place  as  the  type  series  but  is  not  men- 
tioned in  the  original  description. 
Comments.   MCZ  45034  represents  an  immature 
animal  (MacPhee  1987a:  7),  not  an  adult  as  claimed 
in   the   original  description.    MacPhee   (1987a:   9) 
synonyinized  drouhardi  with  Microgale  cowani,  an 
approach  followed  by  Hutterer  (1993:  71),  but  Jen- 
kins et  al.  (1997:  6)  argued  that  M.  droidiardi  is  a 
distinct  species. 

Microgale  parvula  G.  Grandidier,  1934 
Bull.  Mus.  Hist.  Nat.  Paris,  6:  476,  29 
November. 

Holotype.  MCZ  45465.  Body  in  alcohol,  skull  extract- 
ed. Juvenile  male. 


Locality.  Madagascar:  (Antsiranana),  east  coast,  Di- 
ego-Suarez  (=  Antsiranana).  May  1934. 
Collector.  M.  Drouhard. 
Condition.  Alcoholic,  skull  complete. 
Tijpe  Series.  Holotype  only. 

Comments.  MCZ  45465  represents  an  immature 
animal  (MacPhee  1987a:  7),  not  an  adult  as  claimed 
in  the  original  description.  The  holotype  of  Micro- 
gale pulla  Jenkins,  1988  actually  represents  an 
adult  specimen  of  M.  parvula  (Jenkins  et  al.,  1996: 
204).  Considered  a  valid  species  by  Hutterer  (1993: 
71)  and  Nowak  (1999:  190). 


Microgale  prollxacaudata  G.  Grandidier, 

1937 

Bull.  Mus.  Hist.  Nat.  Paris,  9:  348,  25 

November. 

=  Microgale  longicaudata  Thomas,  1882. 

See  MacPhee  (1987a:  9). 


Holotype.  MCZ  45035.  Body  in  alcohol,  skull  extract- 
ed. Juvenile. 

Locality.  Madagascar:  (Antsiranana),  east  coast,  Di- 
ego-Suarez  (=  Antsiranana).  May  1934. 
Collector  M.  Drouhard. 

Condition.   Alcoholic;   skull  partial  (left  tympanic 
bulla  missing).  Mandible  disarticulated. 
Type  Series.  1  paratype;  MCZ  46020,  in  alcohol. 
Comments.  This  specimen  represents  an  immature 
animal  (MacPhee  1987a:  8),  not  an  adult  as  claimed 
in  the  original  description. 


Paramlcrogale  occldentalls  G.  Grandidier 
and  Petit,  1931 

Bull.  Soc.  Zool.  France,  56:  129,  15  June. 
=  Microgale  brevlcaudata  G.  Grandidier, 
1899.  See  MacPhee  (1987a:  9). 


Holotype.  MCZ  45047.  Body  in  alcohol,  skull  extract- 
ed. Juvenile  male. 

Locality.  Madagascar:  (Antananarivo),  northwest  of 
Maintirano,  Andriafeuelo.  1930. 
Collector  M.  A.  de  la  Rue. 
Condition.  Alcoholic,  skull  complete. 
Type  Series.  Holotype  only,  but  see  comments. 
Comments.  P.  occidentalis  is  the  type  species  of  the 
genus    Paramicrogale    G.    Grandidier    and    Petit, 
1931.  This  specimen  represents  an  immature  ani- 
mal (MacPhee  1987a:  7),  not  an  adult  as  claimed 
in  the  original  description.  The  original  description 
mentions  only  a  single  specimen  but  describes  ex- 
tensively the  skeleton  of  P.  occidentalis.  This  is  puz- 
zling, as  the  skeleton  has  not  been  extracted  from 
MCZ  45047  (MacPhee  1987a:  7). 


Type  Specimens  of  Recent  Mammals  •  Helgen  and  McFadden        103 


Genus  SETIFER  Fronep,  1806 

Dasogale  fontoynonti  G.  Grandidier, 

1930a 

Bull.  Acad.  Malgache,  n.  sen,  11:  85  (for 

1928). 

=  Setifer  setosus  {Schreber,  1777).  See 

Poduschka  and  Poduschka  (1982:  261). 


Holotype.   MCZ  45016.   Skin,  skull,  and  postcranial 
skeleton.  Juvenile. 

Localitij.  Madagascar:  east  coast.  1917. 
Collector.    Received  by   G.    Grandidier  from   the 
Academic  Malgache  in  1917. 

Condition.  Sldn  represented  by  a  small  patch  of 
lur.  Skull  partial  (occipital  region  missing).  Postcra- 
nial skeleton  complete,  partially  articulated. 
Tijpe  Series.  1  paratype;  MCZ  45532,  in  alcohol. 
Comments.  Walker  (1975:  110)  commented  that 
"the  only  specimen  known  of  D.  fontoynonti  ...  is 
in  the  Paris  Museum";  actually,  the  only  material 
attributed  to  Dasogale  is  in  the  MCZ,  a  fact  first 
noted  in  publication  by  Poduschka  and  Poduschka 
(1982:  253).  Dasogale  was  often  considered  to  be 
an  extremely  rare  or  recently  extinct  species  until 
Poduschka  and  Poduschka  (p.  261)  and  MacPhee 
(19S7b:  135)  demonstrated  that  the  holotype  is 
probably  a  juvenile  Setifer  .setosus.  D.  fontoynonti 
is  the  type  species  of  the  genus  Dasogale  G.  Gran- 
didier, 1930. 


Order  RODENTIA  Bowdich,  1821 

Family  APLODONTIDAE  Brandt,  1855 

Genus  APLODONTIA  Richardson,  1829 

Aplodontia  californica  columbiana  Taylor, 

1916 

Univ.  California  Publ.  Zool.,  12:  499,  6 

May. 

= Aplodontia  rufa  rainieri  Merriam,  1899. 

See  Dalquest  (1948:  369). 


Holotype.  B1899.  Sldn  and  skull.  Adult  male. 

Locality.     (Canada):    British    Columbia,    Hope, 

Roabs  Ranch.  14  June  1894. 

Collector  W.  C.  Colt.  Original  number  479. 

Condition.  Skin  and  skull  complete. 

Type  Series.  8  paratypes;  B1892-B1898,  B1900;  all 

represented  by  skin   and  skull,   3  females  and  5 

males. 


Family  SCIURIDAE  Fischer  de  Waldheim, 
1817 

Genus  CALLOSCIURUS  Gray,  1867 

Callosciurus  baluensis  medial  is  G.  M. 

Allen  and  Coolidge,  1940 

Bull.  Mus.  Comp.  Zool.,  87:  156,  31 

December. 

Holotype.  MCZ  22265.  Skin  and  skull.  Adult  female. 
Locality.  (Indonesia):  Dutch  Borneo,  (Kalimantan), 
Mount  Tibang  (possibly  =Bukit  Tungun).  1925. 
Collector.  E.  Mjoberg.  Original  number  6. 
Condition.   Skin  complete.  Skull  partial  (right  pa- 
rietal,   right   tympanic   bulla,   and   lachiymal   bro- 
ken— bulla  present).  Mandible  disarticulated. 
Type  Series.  Holotype  only. 

Comments.  Retained  as  a  valid  subspecies  by  Med- 
way  (1977:  90). 

Callosciurus  ferruginous  primus  G.  M. 

Allen  and  Coolidge,  1940 

Bull.  Mus.  Comp.  Zool,  87:  157,  31 

December. 

=  Callosciurus  erythraeus  (Pallas,  1778). 

See  Corbet  and  Hill  (1992:  283). 

Holotype.  MCZ  35352.  Skin  and  skull.  Adult  female. 
Locality.  Siam  (=Thailand):  Mae  Wan  River  near 
Doi,  Mount  Souket  (  =  Saket),  1,500  ft  (458  m).  20 
Febiaiary  1937. 

Collector  J.  A.  Griswold,  Jr.,  Asiatic  Primate  Ex- 
pedition. Original  number  5. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  1  paratype;  MCZ  35353,  skin  and 
skull,  adult  male. 

Sciurus  castaneoventris  haemobaphes  G. 

M.  Allen,  1912c 

Proc.  Biol.  Soc.  Washington,  25:  177,  24 

December. 

=  Callosciurus  erythraeus  haemobaphes 

(G.  M.  Allen,  1912).  See  Hayman  and 

Holt  (1940:  359). 

Holotype.  MCZ  13693.  Skin  and  skull.  Male. 

Locality.   China:  southeastern  Yunnan,  Chih-ping 
(  =  Shiping).  26  Februaiy  1911. 
Collector   Kobayashi  Collection.  Original  number 
46. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Only  the  holotype  is  mentioned  in  the 
original  publication,  but  Allen  had  four  other  spec- 
imens of  these  squirrels  at  the  time  of  description 
(MCZ  13692,  13694-13696;  all  represented  by  skin 
and  skull,  3  males,  1  female). 


104         Bulletin  Museum  of  Comparative  Zoolog^y,  Vol.  157,  No.  2 


Genus  DREMOMYS  Heude,  1 898 

Dremomys  pernyi  flavior  G.  M.  Allen, 

1912c 

Proc.  Biol.  Soc.  Washington,  25:  178,  24 

December. 

Holottjpe.  MCZ  13691.  Skin  and  skull.  Male. 

Locality.    China:   southeastern  Yunnan,   Mongtz 
(  =  Mengzi).  1911. 

Collector.   Kobayashi  Collection.  Original  number 
6/8. 

Condition.   Skin  complete.  Skull  partial  (squamo- 
sals  missing,   supraoccipital   damaged,   right  jugal 
missing,    palatine    missing,    tympanic    bulla    dam- 
aged). 
Type  Series.  Holoty]^)e  only. 

Dremomys  senex  G.  M.  Allen,  1912b 

Mem.  Mus.  Comp.  ZooL,  40:  229, 

August. 

=  Dremomys  pernyi  senex  G.  M.  Allen, 

1912.  See  Hayman  and  Holt  (1940:  382). 

Holotype.  MCZ  7582.  Skin  and  skull.  Adult  female. 
Locality.    China:    Hupeh   (  =  Hubei),   Ichanghsien, 
Nantou.  5  Februaiy  1909. 

Collector  W.  R.  Zappey.  Original  number  .373. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  1  paraty^je;  MCZ  7583,  skin  and  skull, 
adult  male. 

Genus  FUNISCIURUS  Trouessart,  1880 

Funisciurus  pyrrhopus  victoriae  G.  M. 

Allen  and  Loveridge,  1942 

Bull.  Mus.  Comp.  Zool.,  8:  180, 

February. 

=  Funisciurus  pyrriiopus  al<l<a  De  Winton, 

1899.  See  Amtmann  (1975:  8). 

Holotype.  MCZ  39199.  Skin  and  skull.  Adult  male. 
Locality.    Uganda:   Toro,    Kibale   Forest,   4,200   ft 
(1,281  m).  16  December  1938. 
Collector  A.  Loveridge. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 
Type  Series.  Holotype  only. 

Genus  GLAUCOIVIYS  Thomas,  1908 

Sciuropterus  alpinus  bangsi  Rhoads,  1897 
Proc.  Acad.  Nat.  Sci.  Philadelphia,  1897, 
p.  321  (footnote),  July. 
^Glaucomys  sabrinus  bangsi  (Rhoads, 
1897).  See  A.  H.  Howell  (1918:  38). 

Holotype.  B6959.  Sldn  and  skull.  Adult  male. 

Locality.    (United   States):    Idaho,    Idalio   County. 
March  8,  1897. 


Collector  Harbison  and  Bargamin. 
Condition.  Skin  and  skull  complete. 
Type   Series.    1   paratype;   B6960,   skin   and  skull, 
adult  male. 

Comments.  G.  s.  bangsi  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  450). 

Sciuropterus  alpinus  lascivus  Bangs, 

1899J 

Proc.  New  England  Zool.  Club,  1:  69,  31 

July. 

=  Glaucomys  sabrinus  lascivus  (Bangs, 

1899).  See  A.  H.  Howell  (1918:  55). 

Holotype.  B9186.  Skin  and  skull.  Adult  female. 
Locality.    (United   States):   California,   El   Dorado 
County,  Tallac.  28  August  1898. 
Collector  W.  VV.  Price  and  P.  O.  Simons.  Original 
number  1722. 

Condition.  Sldn  and  skull  complete. 
Type  Series.  2  paratypes,  B9187,  skin  and  skull,  fe- 
male; B9188,  sldn  and  skrdl,  female. 
Comments.   G.  s.  la.scivns  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  451). 

Sciuropterus  sabrinus  makkovikensis 

Sornborger,  1900 

Ottawa  Nat.,  14:  48,  6  June. 

=  Glaucomys  sabrinus  makkovikensis 

(Sornborger,  1900).  See  A.  H.  Howell 

(1918:  34). 

Syntypes.  MCZ  10476:  Skin  and  skull.  Adult.  MCZ 
10477:  Skin  and  skull.  Adult.  MCZ  10478:  Skin  and 
skull.  Adult. 

Locality.  (Canada):  Labrador  Peninsula,  Makkovik. 
1899. 

Collector.  W.  W.  Perrett.  Original  number  1540. 
Condition.  MCZ  10476:  Skin  complete.  Skull  par- 
tial (base  of  skull  missing).  Mandible  disarticulated. 
MCZ  10477:  Skin  partial  (tail  broken  but  present). 
Skull  partial  (base  of  skull  missing).  Mandible  dis- 
articulated. MCZ  10478;  Skin  partial  (missing  left 
hind  foot  and  tail).  Skull  partial  (occipital  chipped). 
Mandible  disarticulated. 
Type  Series.  3  syntypes  only. 

Comments.  Sornborger's  original  description  was 
based  on  three  specimens,  original  numbers  1540, 
1541,  and  1542,  now  MCZ  10476,  10477,  10478, 
respectively.  G.  .$.  makkovikensis  was  retained  as  a 
valid  subspecies  by  Hall  (1981:  453). 

Sciuropterus  silus  Bangs,  1896J 

Proc.  Biol.  Soc.  Washington,  10:  163,  28 

December. 

=  Glaucomys  volans  volans  (Linnaeus, 

1758).  See  A.  H.  Howell  (1918:  20). 

Holotype.  B4931.  Sldn  and  skull.  Adult  male. 

Locality.  (United  States):  West  Virginia,  Greenbri- 


Type  Specimens  of  Recent  Mammals  •  Helpen  and  McFodden 


105 


er  County,  White   Sulphur  Springs,  top  of  Katis 
Mtn.  3,200  ft  (976  m).  2  September  1895. 
Collector.  T.  Surber.  Original  number  19. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 
Type  Series.  Holotype  only. 

Sciuropterus  volans  querceti  Bangs,  1 896j 

Proc.  Biol.  Soc.  Washington,  10:  166,  28 

December. 

=  Glaucomys  volans  querceti  (Bangs, 

1896).  See  A.  H.  Howell  (1918:  26). 

Holotype.  B2451.  Skin  and  skull.  Adult  female. 
Locality.   (United  States):  Florida,  Citrus  County, 
Citronelle.  17  September  1894. 
Collector  F.  L.  Small.  Original  number  1363. 
Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.   2  paratypes;  B2452,  skin  and  skull, 
adult  male;  B2453,  skin  and  skull,  adult  female. 
Comments.  G.  v.  querceti  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  449). 

Genus  HYLOPETES  Thomas,  1 908 

Pteromys  (Hylopetes)  alboniger  orinus  G. 

M.  Allen,  1940 

The  Mammals  of  China  and  Mongolia, 

Natural  Histoiy  of  Central  Asia,  11:  723, 

3  September. 

=  Hylopetes  alboniger  alboniger  (Hodgson, 

1836).  See  Ellerman  and  Morrison-Scott 

(1951:  469). 

Holotype.  MCZ  28086.  Skin  and  skull.  Adult  female. 
Locality.    China:    Yunnan,    Likiang    Range    (  =  Li- 
jiang),  7,800  ft  (2,379  m).  December  1931. 
Collector  J.  F  Rock. 

Condition.  Skin  complete.  Skull  partial  (most  of 
right  tympantic  bulla  missing,  supraoccipital  dam- 
aged). Mandible  disarticulated. 
Type  Series.  Allen  examined  10  specimens  in  ad- 
dition to  the  type,  including  4  specimens  from  the 
BMNH,  3  from  AMNH,  and  MCZ  28087,  skin  and 
skull,  an  unsexed  adult. 

Pteromys  phayrei  anchises  G.  M.  Allen 

and  Coolldge,  1940 

Bull.  Mus.  Comp.  Zool.,  87:  153,  31 

December. 

=  Hylopetes  phayrei  anchises  (G.  M.  Allen 

and  Coolldge,  1940).  See  Ellerman  and 

Morrison-Scott  (1951:  469). 

Holotype.  MCZ  35776.  Skin  and  skull.  Adult  male. 
Locality.   Siam  (=Thailand):  (Chiang  Mai)  Mount 


Angka  (  =  Doi  Inthanon),  4,300  ft  (1,312  m).  27 
February  1937. 

Collector.  J.  A.  Griswold,  Jr.,  Asiatic  Primate  Ex- 
pedition. Original  number  24. 

Condition.  Skin  and  skvdl  complete.  Mandible  dis- 
articulated. 

Type  Series.  3  paratypes;  MCZ  35775,  skin  and 
sk-ull,  adult  female;  MCZ  35777,  skin  and  skull, 
adult  male;  MCZ  35778,  skin  and  skull,  subadult 
male. 

Genus  MARMOTA  Blumenbach,  1779 

Arctomys  flaviventer  avarus  Bangs,  1899] 

Proc.  New  England  Zool.  Club,  1:  68,  31 

July. 

=  Marmota  flaviventris  avara  (Bangs, 

1899).  See  A.  H.  Howell  (1915:  41). 

Holotype.  B7299.  Skin  and  skull.  Juvenile  female. 
Locality.    (Canada):   British  Columbia,  Okanagan. 
17  July  1897. 

Collector  A.  C.  Brooks.  Original  number  969. 
Condition.  Skin  and  skadl  complete. 
Type  Series.  2  paratypes;  B7298,  skin  and  skull,  ju- 
venile male;  B7300,  skin  and  skull,  juvenile  female. 
Comments.  M.  f.  avara  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  371). 

Arctomys  ignavus  Bangs,  1899d 

Proc.  New  England  Zool.  Club,  1:  13,  28 

February. 

=  Marmota  monax  ignava  (Bangs,  1899). 

See  A.  H.  Howell  (1915:  29). 

Holotype.  B7971.  Skin  and  skull.  Adult  male. 

Locality.  (Canada):  Labrador  Peninsula,  Black  Bay. 
13  July  1898. 
Collector.  E.  Doane. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  5  paratypes;  B7968-B7970,  B7972, 
B7973  (juvenile);  all  represented  by  skin  and  skull, 
4  females  and  1  male. 

Comments.  M.  m.  ignava  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  370)  and  Kwiecinski 
(1998:  1). 

Genus  MICROSCIURUS  J.  A.  Allen,  1895 

Sciurus  (Microsciurus)  browni  Bangs, 

1902b 

Bull.  Mus.  Comp.  Zool.,  39:24,  April. 

=  Microsciurus  alfari  browni  (Bangs,  1902). 

See  J.  A.  Allen  (1914:151). 

Holotype.  MCZ  10404.  Skin  and  skull.  Adult  male. 
Locality.    Panama:   Chiriqui,   Bogaba,  600  ft  (183 
m).  15  July  1901. 


106         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Collector.  W.  W.  Browii,  Jr.  Original  number  631. 
Condition.  Sldn  and  skull  complete. 
Type   Series.   4  paratypes;   MCZ   10405,  adult  fe- 
male,   10406,   adult  female,    10407,   adult  female, 
10408,  juvenile  male;  all  represented  by  sldn  and 
skull. 

Comments.  M.  a.  broivni  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  439). 

Genus  PARAXERUS  Forsyth  Major,  1893 

Aethosciurus  byatti  laetus  G.  M.  Allen  and 

Loverldge,  1933 

Bull.  Mus.  Comp.  Zool,  75:  96, 

Februaiy. 

=  Paraxerus  vexillarius  byatti  (Kershaw, 

1923).  See  Amtmann  (1975:  11). 

Holotype.  MCZ  26198.  Skin  and  skull.  Adult  male. 
Locality.  Tanganyika  Territory  (=Tanzania):  north 
end  of  Lake  Nyasa,  Ukinga  Mountains,  Madehani, 
7,000  ft  (2,135  m).  22  Februaiy  1930. 
Collector.  A.  Love  ridge. 
Condition.  Sldn  and  skull  complete. 
Type    Series.    9   paratypes;    MCZ   26196,    26197, 
26199-26202,    26204-26206;    all    represented   by 
skin  and  skull,  4  females  and  5  males. 

Genus  SCIUROTAMIAS  M\\\er,  1901 

Sciurotamias  davidanus  thayeri  G.  M. 

Allen,  1912b 

Mem.  Mus.  Comp.  Zool.,  40:  231, 

August. 

=  Sciurotamias  davidianus  consobrinus 

(Milne-Edwards,  1868).  See  Moore  and 

Tate  (1965:  308). 

Holotype.  MCZ  8008.  Skin  and  skull.  Adult  male. 
Locality.    China:    western    Szechwan    (  =  Sichuan), 
Washan  (=Wushan),  6,000  ft  (1,830  m).   17  May 
1908. 

Collector  W.  R.  Zappey.  Original  number  163. 
Condition.  Sldn  complete.  Skull  partial  (basioccip- 
ital  and  left  maxilla  damaged).  Mandible  disartic- 
ulated. 

Type  Series.  Holotype  only. 

Comments.  G.  M.  Allen  (1912b:  231)  noted  in  the 
original  description  of  thayeri  that  "unfortunately, 
tlie  skull  of  the  type  was  lost."  The  sldn  and  skull 
have  been  revmited  subsequently. 

Genus  SC/L/RL/S  Linnaeus,  1758 

Sciurus  (Guerlinguetus)  aestuans 
cfiiriquensis  Bangs,  1902b 
Bull.  Mus.  Comp.  Zool.,  39:  22,  April. 
=  Sciurus  granatensis  chiriquensis  Bangs, 
1902.  See  Hershkovitz  (1947:  7). 

Holotype.  MCZ  10044.  Skin  and  skull.  Adult  male. 
Locality.  Panama:  Chiriqui,  Divala.  18  November 
1900. 


Collector.  W.  W.  Brown,  Jr.  Original  number  10. 
Condition.  Skin  and  skull  complete. 
Type  Series.  There  is  a  large  series  of  paratypes  in 
the  MCZ. 

Comments.  S.  g.  chiriquensis  was  retained  as  a  val- 
id subspecies  by  Hall  (1981:  437)  and  Nitikman 
(1985:  1). 

Sciurus  carolinensis  extimus  Bangs,  1896j 
Proc.  Biol.  Soe.  Washington,  10:  158,  28 
December. 

Holotype.  B4519.  Sldn  and  skull.  Adult  female. 

Locality.    (United  States):   Florida,  Dade  County, 

Miami.  12  March  1895. 

Collector.  L.  Brownell.  Original  number  59. 

Condition.  Skin  complete  (with  bald  spots  on  ven- 

trum).  Skull  complete. 

Type  Series.   7  paratypes;  B3406,  B4517,  B4518, 

(all  represented  by  skin  and  skull),  B4520-B4523 

(skins  only);  4  females  and  3  males. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 

(1981:  417)  and  Koprowskd  (1994:  1). 

Sciurus  carolinensis  var.  yucatanensis  J. 

A.  Allen,  1877 

In  Coues  and  J.  A.  Allen,  Monogr.  N. 

Amer.  Rodentia,  U.S.  Geol.  Geograph. 

Suivey  Terr.,  Rep.  11:  705,  August. 

=  Sciurus  yucatanensis  J.  A.  Allen,  1877. 

See  Elliot  (1896:  80). 

Syntype.  MCZ  5398.  Skin.  Adult  male. 

Locality.  (Mexico):  Yucatan,  Merida.  March  1865. 
Collector.    A.   Schott.   Original   number  228.   For- 
merly USNM  8502. 
Condition.  Sldn  complete. 

Type  Series.  J.  A.  Allen  referred  to  "four  specimens 
of  this  variety  before  me"  in  the  original  descrip- 
tion (1877:  705).  Three  specimens  are  mentioned 
by  number:  USNM  8502  (now  MCZ  5398),  8503, 
and  8505.  A  juvenile  referred  to  in  the  description 
(but  not  by  number)  corresponds  to  USNM  8504. 
Comments.  S.  yucatanensis  was  considered  a  valid 
species  by  Hoffman  et  al.  (1993:  443)  and  Nowak 
(1999:  1265).  The  syntype  of  yucatanensis  in  the 
MCZ  was  received  from  the  USNM  in  March 
1877.  Poole  and  Schantz  (1942:  554)  state  that 
USNM  8505  is  no  longer  in  the  USNM. 

Sciurus  ludovicianus  vicinus  Bangs,  1896j 

Proc.  Biol.  Soc.  Washington,  10:  150,  28 

December. 

=  Sciurus  niger  vulpinus  Gmelin,  1896. 

See  Bark-alow  (1954:  25). 

Holotype.  B5215.  Skin  and  skull.  Adult  female. 
Locality.  (United  States):  West  Virginia,  Greenbri- 


Type  Specimens  of  Recent  Mammals  •  Helgen  and  McFodden        107 


ei"    County,    White    Sulphur    Springs.    29   Januaiy 

1896. 

Collector.  T.  Surber.  Original  number  55. 

Condition.  Sldn  and  skull  complete. 

Type  Series.  There  is  a  series  of  paratypes  in  the 

MCZ. 

Sciurus  nesaeus  G.  M.  Allen,  1902b 

Proc.  Biol.  Soc.  Washington,  15:  93,  25 

April. 

=  Sciurus  granatensis  nesaeus  G.  M. 

Allen,  1902.  See  Hershko\atz  (1947:  37). 

Holotijpe.  MCZ  10744.  Skin.  Adult  female. 

Locality.    Venezuela:    (Nueva   Esparta),   Margarita 

Island,  El  Valle.  8  July  1901. 

Collector.  A.  H.  Clark.  Original  number  619. 

Condition.  Skin  complete.  The  lower  incisors  are 

present  in  the  sldn. 

Type  Series.  Holotype  only. 

Comments.   S.  g.   nesaeus  was  retained  as  a  valid 

subspecies  by  Cabrera  (1961:  367)  and  Nitikman 

(1985:  1). 

Sciurus  variabilis  morulus  Bangs,  1 900d 

Proc.  New  England  Zool.  Club,  2:  43,  20 

September. 

=  Sciurus  granatensis  morulus  Bangs, 

1900.  See  Miller  and  Kellogg  (1955: 

257). 

Holotype.  B8420.  Skin  and  skull.  Adult  female. 

Locality.  Panama:  Canal  Zone,  Loma  del  Leon.  13 

March  'l900. 

Collector  W.  W.  Brown,  Jr.  Original  number  5. 

Condition.   Skin  complete.  Skull  slightly  damaged 

(left  bulla  broken). 

Type  Series.   5  paratypes;  B8418,  B8419,  B8421- 

B8423  (juvenile);  all  represented  by  skin  and  skull, 

3  females  and  2  males. 

Comments.   S.  g.  mondus  was  retained  as  a  valid 

subspecies    by    Hall    (1981:    437)    and    Nitikman 

(1985:  1). 

Sciurus  variabilis  saltuensis  Bangs,  1898o 

Proc.  Biol.  Soc.  \\'ashington,  12:  185,  16 

November. 

=  Sciurus  granatensis  saltuensis  Bangs, 

1898.  See  Hershkovitz  (1947:  15). 

Holotype.  B8144.  Skin  and  skull.  Adult  female. 

Locality.    Colombia:    Magdalena,    Santa    Marta 

Mountains,  Pueblo  Viejo,  8,000  ft  (2,440  m).  26 

March  1898. 

Collector.  W.  W.  Brown,  Jr.  Original  number  112. 

Condition.  Sldn  and  skull  complete. 

Type  Series.   2  paratypes;  B8145,  skin  and  skull, 

adult  male;  B8244,  skin  and  skull,  adult  female. 

Comments.  S.  g.  saltuensis  was  retained  as  a  valid 


subspecies  by  Cabrera  (1961:  368)  and  Nitikman 

(1985:  1). 

Genus  SPERMOPHILUS  F.  Cuvier,  1825 

Citellus  obscurus  siccus  G.  M.  Allen,  1925 
Amer.  Mns.  Novitates,  163:  3,  2  April. 
=  Spermophilus  alashanicus  Buchner, 
1888.  See  Hoffmann  et  al.  (1993:  444). 

Holotype.  MCZ  19924.  Skin  and  skull.  Adult  female. 
Locality.  China:  Shansi  (  =  Slienxi),  10  miles  (16.1 
km)  west  of  Taiyuanfu.  August  1921. 
Collector.  F.  R.  Wulsin.  Original  number  146. 
Condition.   Skin  complete.  Skull  partial  (parietals 
and   basiooccipital    missing,    right   tympanic   bulla 
broken  but  present).  Mandible  disarticulated. 
Ti/pe  Series.  Holotype  only. 

Spermophilus  armatus  Kennicott,  1863 
Proc.  Acad.  Nat.  Sci.  Philadelphia,  15: 
158,  June. 

Sipitypes.  MCZ  297:  Skull.  Male.  Collected  11  April 
'  1858.  Original  number  167,  formerly  USNM  4799. 
MCZ  4793:  Skin  and  skull.  Male.  Collected  2  April 
1858.  Original  number  140,  formerly  USNM  3478 
(3373).  MCZ  4790:  Skin.  Female.  Collected  26 
May  1858.  Original  number  455,  formerly  USNM 
3470  (3470).  MCZ  4794:  Skull  only  (skin  spoiled  in 
repreparation,  discarded).  Male.  Collected  April 
14,  1858.  Original  number  215,  formerly  USNM 
3474  (3373). 

Locality.  (United  States):  Utah  (now  Wyoming), 
(Uinta  County),  near  Fort  Bridger,  foothills  of  the 
Uinta  Mountains,  Camp  Scott. 
Collector  C.  Drexler. 
Type  Series.  See  comments  below. 
Comments.  Considered  a  valid  species  by  Hoff- 
mann et  al.  (1993:  444)  and  Nowak  (1999:  1254). 
Kennicott  does  not  designate  a  type  in  the  original 
descidption;  thus,  "all  the  specimens  from  Fort 
Bridger  collected  by  C.  Drexler  and  in  the  collec- 
tions [of  the  USNM]  prior  to  1863  are  evidently 
cotypes  [  =  syntypes]  of  this  species"  (Lyon  and  Os- 
good 1909:  163).  The  syntypes  of  annatus  in  the 
MCZ  were  received  from  the  USNM  in  January 
1874.  These  specimens  possess  USNM  labels  that 
bear  numbers  in  discrepancy  with  the  USNM  cat- 
alogue; the  numbers  listed  above  are  taken  from 
the  catalogue  of  the  USNM,  followed  in  parenthe- 
ses by  the  number  on  the  specimen  label.  A  com- 
plete list  of  syntypes,  most  of  the  remainder  of 
which  are  in  the  USNM,  can  be  found  in  Poole 
and  Schantz  (1942:  504-505).  The  skin  of  MCZ 
297,  which  formerly  bore  tlie  number  USNM 
3472,  is  not  to  be  found  in  the  collections  of  either 


108         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Spermophilus  elegans  Kennicott,  1863 
Proc.  Acad.  Nat.  Sci.  Philadelphia,  15: 
158,  June. 

Syntijpes.  MCZ  4791:  Skin.  Female.  Collected  11 
April  1858.  Original  number  168.  Formerly  USNM 
3468.  MCZ  4792:  Skin.  Male.  Collected  11  April 
1858.  Original  number  166.  Formerly  USNM 
3473. 

Locality.    (United  States):   Utah   (now  Wyoming), 
(Uinta  County),  Fort  Bridger 
Collector.  C.  Dre.-der. 

Condition.  Following  their  receipt,  these  speci- 
mens were  spoiled  in  repreparation  and  discarded. 
Type  Series.  See  comments  below. 
Comments.  These  specimens  are  no  longer  in  ex- 
istence. Kennicott  does  not  designate  a  type  in  the 
original  description;  thus,  "all  the  specimens  from 
Fort  Bridger  collected  by  C.  Drexler  and  in  the 
collections  [of  the  USNM]  prior  to  1863  are  evi- 
dently cotypes  [  =  syntypes]  of  this  species"  (Lyon 
and  Osgood  1909:  163).  These  syntypes  oi  elegans 
were  received  from  the  USNM  in  January  1874.  A 
complete  list  of  syntypes  can  be  found  in  Lyon  and 
Osgood  (1909:  166).  G.  M.  Allen  (1931:  251)  er- 
roneously included  MCZ  4791  in  a  list  of  the  type 
series  of  Spennophihis  annatus.  S.  elegans  is  con- 
sidered a  valid  species  by  Hoffmann  et  al.  (199.3: 
446)  and  Nowak  (1999:  1254). 

Spermophilus  (Ictidomys)  tridecemlineatus 

badius  Bangs,  1899c 

Proc.  New  England  Zool.  Club,  1:  1,  8 

Februaiy. 

=  Spermophilus  tridecemlineatus  texensis 

Merriam,  1898.  See  Hall  and  Kelson 

(1959:  347). 

Holotype.  B1682.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Missouri  (Vernon  Coun- 
ty), Stotesbury.  17  April  1894. 
Collector.  T.  Surber.  Original  number  81. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.   2  paratypes;   B1683,  skin  and  skull, 
adult  male;  B5609,  skin  and  skull,  adult  male. 
Comments.  A.  H.  Howell  (1938:  110)  first  synon- 
yinized  badius  with  texensis  under  die  genus  Ci- 
tellus. 

Genus  SYNTHEOSCIURUS  Bangs,  1902 

Syntheosciurus  brochus  Bangs,  1 902b 
Bull.  Mus.  Comp.  Zool.,  39:  25,  April. 

Holotype.  MCZ  10402.  Sldn  and  skull.  Adult  male. 
Locality.   Panama:  Chiriqui,  Boquete  (  =  Bajo  Bo- 
quete),  7,000  ft  (2.135  m).  30  April  1901. 
Collector  W.  W.  Brown,  Jr  Original  number  415. 


Condition.  Skin  complete.  Skull  partial  (left  supra- 
occipital  damaged) 

Type  Series.  1  paratype;  MCZ  10403,  sldn  and 
skull,  adult  female. 

Comments.  Type  species  of  the  genus  Syntheo.sciu- 
nis  Bangs,  1902.  Considered  a  valid  species  by 
Hoffman  et  al.  (1993:  452)  and  Nowak  (1999: 
1268).  Following  a  discussion  with  a  collector  em- 
ployed by  W.  W.  Brown,  Jr.,  and  a  biological  survey 
of  Boquete,  Enders  (1953b:  509)  recommended 
that  the  type  locality  be  considered  "the  Cordillera 
about  8  miles  [12.9  km]  north  of  Boquete  and  not 
at  Boquete  which  is  on  the  lower  slopes  of  El  Vol- 
can  de  Chiriqui." 

Genus  LAMMS  llliger,  1811 

Tamias  cooperi  Baird,  1855 

Proc.  Acad.  Nat.  Sci.  Philadelphia,  7: 

334,  24  April  24. 

=  Tamias  townsendii  cooperi  Baird,  1855. 

See  Baird  (1858:  737). 

Syntype.  MCZ  4754.  Skin  only.  Adult,  unsexed. 
Locality.    (United    States):    Washington    (Skagit 
County),    Clickitat    (  =  Klickitat)    Pass,    Cascade 
Mountains,  4,500  ft  (1,373  m).  July  1853.  See  com- 
ments. 

Collector  J.  G.  Cooper.  Formerly  USNM  211/ 
1182. 

Condition.  Sldn  complete. 

Type  Series.  The  other  syntype  is  USNM  212/1183, 
skin  and  skull,  unsexed  adult. 

Comments.  No  type  is  designated  in  the  original 
description,  but  Baird  subsequently  ex-plained  that 
the  two  specimens  listed  above  were  those  on 
which  cooperi  was  based  (1858:  301).  Cooper 
(1869:  .531)  emended  the  type  locality.  The  syntype 
of  cooperi  in  the  MCZ  was  received  from  the 
USNM  in  Januarv'  1874. 

Tamias  dorsalis  Baird,  1855 

Proc.  Acad.  Nat.  Sci.  Philadelphia,  7: 

332,  24  April. 

=  Tamias  dorsalis  dorsalis  Baird,  1855. 

See  Hayman  and  Holt  (1940:  435). 

Syntype.  MCZ  4759.  Skin  and  mandible. 

Locality.  (United  States):  New  Me.xico  (Grant 
County),  "Fort  Webster,  Coppermines  of  die  Mim- 
bres"  (near  present  Georgetown).  32°47'N, 
108°41'W.  1851.  See  A.  H.  Howell  (1929:  131). 
Collector  J.  H.  Clark.  Formerly  USNM  119/3151. 
Condition.  Skin  complete.  Mandible  partial  (pos- 
terior of  right  mandibular  ramus  broken  off  and 
missing). 

Type  Series.  The  other  syntype  is  USNM  120,  skull 
within  skin. 

Comments.  No  type  is  designated  in  the  original 
description,  but  Baird  subsequently  listed  the  two 


Type  Specimens  of  Recent  Mammals  •  Helaen  and  McFadden 


109 


specimens  above  as  those  on  which  dorsalis  was 
based  (1858:  300).  The  syntype  of  dorsalis  in  the 
MCZ  was  received  from  the  USNM  in  January 
1874. 

Tamias  quadrivittatus  neglectus  J.  A. 

Allen,  1890 

Bull.  Amer.  Mus.  Nat.  Hist.,  3:  106, 

June. 

=  Tamias  minimus  neglectus  (J.  A.  Allen, 

1890).  See  Hayman  and  Holt  (1940: 

430). 

Holotype.  MCZ  1575.  Sldn  and  cranium. 

Locality.  (Canada):  (Ontario),  eastern  end  of  Lake 
Superior  (near  mouth  of  Montreal  River).  5  July 
1848. 

Collector.  L.  Agassiz. 

Condition.  Sldn  complete.  Cranium  partial  (zygo- 
matic arches,  parietals,  left  tympanic  bulla,  and 
mandible  missing). 

Type  Series.  The  description  mentions  6  paratypes, 
including  MCZ  1567,  skin  and  skull;  and  4  others 
at  USNM. 

Comments.  Following  A.  H.  Howell  (1929:  54), 
Hayman  and  Holt  (1940:  430)  considered  T.  m. 
neglectus  a  synonym  of  T.  m.  borealis.  However, 
neglectus  was  retained  as  a  valid  subspecies  of  min- 
imus by  Hall  (1981:  346)  under  the  genus  Euta- 
mias. 

Tamias  striatus  venustus  Bangs,  1 896h 
Proc.  Biol.  Soc.  Washington,  10:  137,  28 
December. 

Holotype.  B5478.  Skin  and  skull.  Adult  male. 

Locality.      (United     States):      Indian     Territory 

(  =  Oklahoma)  (Adair  County),  Stilwell.  13  August 

1896, 

Collector  T.  Surber.  Original  number  63. 

Condition.  Skin  complete.  Skull  partial  (condyle  of 

left  mandibular  ramus  inissing). 

Type  Series.   2  paratypes;  B5479,  skin  and  skull, 

adult  female;  B5605,  sldn  and  skull,  adult  male. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 

(1981:  340). 

Genus  r>4/WMSC/L/f?L/S  Trouessart,  1880 

Sciurus  iiudsonicus  gymnicus  Bangs, 

18991 

Proc.  New  England  Zool.  Club,  1:  28,  31 

March. 

=  Tamiasciurus  Iiudsonicus  gymnicus 

(Bangs,  1899).  See  Osgood  (1938:  438). 

Holotype.  B4914.  Skin  and  skull.  Adult  female. 
Locality.      (United     States):     Maine,     Piscataquis 


County,  Greenville,  near  Moosehead  Lake.  1  De- 
cember 1895. 

Collector  C.  H.  Goldthwaithe.  Original  number  2. 
Condition.  Skin  and  skull  complete. 
Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  T.  h.  gymnicus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  442)  and  Steele  (1998: 
1). 

Sciurus  hudsonicus  loquax  Bangs,  1 896j 

Proc.  Biol.  Soc.  Washington,  10:  161,  28 

December. 

=  Tamiasciurus  hudsonicus  loquax 

(Bangs,  1896).  See  A.  H.  Howell  (1936: 

1). 

Holotype.  B4270.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Connecticut,  New  Lon- 
don County,  Liberty  Hill.  24  December  1895. 
Collector.  O.  Bangs.  Original  number  3. 
Condition.  Skin  and  skull  complete. 
Type  Series.  There  is  a  large  series  of  paratypes  in 
the  MCZ. 

Comments.  T.  h.  loquax  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  443)  and  Steele  (1998:  1). 

Sciurus  hudsonicus  orarius  Bangs,  18971 

Proc.  Biol.  Soc.  Washington,  11:  281,  30 

December. 

=  Tamiasciurus  douglasii  mollipilosus 

(Audubon  and  Bachman,  1841).  See 

Hayman  and  Holt  (1940:  347). 

Holotype.  B4978.  Skin  and  skull.  Adult  female. 

Locality.    (United   States):   California,   Mendocino 
County,  Philo.  9  December  1895. 
Collector.  C.  A.  Allen.  Original  number  887. 
Condition.  Skin  complete.  Skull  partial  (right  tym- 
panic bulla  broken). 

Type  Series.  13  paratypes;  B4832,  B4979-B4989, 
B5462;  all  represented  by  sldn  and  skull,  9  feinales 
and  4  males. 

Family  CASTORIDAE  Hemprich,  1820 

Genus  C/^SrOR  Linnaeus,  1758 

Castor  caecator  Bangs,  1 91 3 
Bull.  Mus.  Comp.  Zool.,  54:  513,  July. 
=  Castor  canadensis  caecator  Bangs, 
1913.  See  G.  M.  Allen  (1942:  62). 

Holotype.  B6979.  Skull.  Adult  male. 

Locality.    (Canada):   Newfoundland,  near  Bay  St. 
George.  1896. 
Collector.  E.  Doane. 

Condition.  Skull  complete.  Mandible  disarticulat- 
ed. 


110         Bulletin  Museum  of  Comparative  Zoologi/,  Vol.  157,  No.  2 


Type  Series.  Holotype  only. 

Comments.  C.  c.  caecator  was  retained  as  a  valid 

subspecies  by  Hall  (1981:  602). 

Family  GEOMYIDAE  Bonaparte,  1845 

Genus  GEOMVS  Rafinesque,  1817 

Geomys  colonus  Bangs,  1898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  178,  15 

March. 

=  Geomys  pinetis  pinetis  Rafinesque, 

1817.  See  Williams  and  Genoways  (1980: 

444). 

Holotijpe.  B5001.  Skin  and  skull.  Advilt  male. 

Locality.  (United  States):  Georgia,  Camden  Coun- 
ty, Arnot  Plantation,  about  4  miles  (6.4  km)  west 
of  St.  Marys.  21  March  1896, 
Collector.  O.  Bangs.  Original  number  8. 
Condition.  Sldn  and  skull  complete. 
Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  Hall  (1981:  505)  recognized  Geomys 
colonus  as  a  valid  species. 

Geomys  cumberlandius  Bangs,  1898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  180,  15 

March. 

=  Geomys  pinetis  pinetis  Rafinesque, 

1817.  See  Wihiams  and  Genoways  (1980: 

444). 

Holotype.  B5016.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Georgia,  Camden  Coun- 
ty, Cumberland  Island,  Stafford  Place.  17  April 
1896. 

Collector  O.  Bangs.  Original  number  1. 
Condition.  Sldn  and  skull  complete. 
Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  In  the  original  description.  Bangs  lists 
B5015  as  the  type  and  gives  its  data.  The  data  given 
lit  B5016,  not  B5015.  B5016  has  "Type"  written 
after  the  entiy  in  Bangs'  catalogue  and  should  be 
considered  the  holotype,  rather  than  B5015.  Hall 
(1981:  505)  and  Laerm  (1981:  150)  supported  the 
specific  status  of  Geomys  cumherlandius. 

Geomys  floridanus  austrinus  Bangs, 

1898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  177,  15 

March. 

=  Geomys  pinetis  pinetis  Rafinesque, 

1817.  See  WilHams  and  Genoways  (1980: 

444). 

Holotype.  B6983.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Florida,  Pinellas  County, 
Belleair.  3  August  1897. 


Collector.  W.  S.  Dickinson. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 
Type  Series.  1  paratype,  in  the  USNM. 

Geomys  tuza  gfof// Sherman,  1944 
Proc.  New  England  Zool.  Club,  23:  38, 
30  August. 

=  Geomys  pinetis  goffi  Slierman,  1944. 
See  Harper  (1952:  37). 

Holotype.  B7222.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Florida,  Brevard  County, 

Eau  Gallie.  18  March  1897. 

Collector.  O.  Bangs.  Original  number  1. 

Condition.  Sldn  and  skull  complete. 

Type  Series.   12  paratypes;  B7212-B7217,  B7219- 

B7221;  all  represented  by  sldn  and  skull,  5  females 

and  7  males. 

Comments.   Williams   and  Genoways   (1980:   444) 

synonymized  gojfi  with  G.  pinetis  pinetis;  however. 

Hall  (1981:  504)  maintained  G,  p.  goffi  as  a  valid 

subspecies.  The  lUCN  designates  goffi  as  extinct. 

Genus  ORTHOGEOMYS  Merriam,  1895 

IVIacrogeomys  cavator  Bangs,  1902b 
Bull.  Mus.  Conip.  Zool.,  39:  42,  April. 
=  Orttiogeomys  cavator  cavator  (Bangs, 
1902).  See  Russell  (1968a:  532). 

Holotijpe.  MCZ  10381.  Skin  and  sk-ull.  Adult  male. 
Locality.   Panama:  Chiriqui,  Boquete  (  =  Bajo  Bo- 
quete),  4,800  ft  (1,464  m).  9  March  1901. 
Collector  W.  W.  Brown,  Jr.  Original  number  212. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articvdated. 

Type  Series.  There  is  a  large  series  of  paratypes  in 
the  MCZ. 

Comments.  O.  cavator  was  considered  a  valid  spe- 
cies by  Patton  (1993:  470)  and  Nowak  (1999:  1314). 

Macrogeomys  pansa  Bangs,  1902b 
Bull.  Mus.  Comp.  Zool.,  39:  44,  April. 
=  Ortiiogeomys  cavator  pansa  (Bangs, 
1902).  See  Russell  (1968a:  532). 

Holotype.  MCZ  10364.  Skin  and  skull.  Adult  female. 
Locality.  Panama:  Chiriqui,  Bogaba,  600  ft  (183 
m).  6  July  1901. 

Collector  W.  W.  Brown,  Jr  Original  number  567. 
Condition.  Skin  complete.  Skull  partial  (right  tym- 
panic bulla  missing). 

Type  Series.  There  are  7  paratypes;  MCZ  10362— 
10363,  10365-10369,  4  females  and  3  males,  all 
represented  by  skin  and  skull.  MCZ  10362  and 
10366  are  now  in  the  FMNH,  and  MCZ  10365  is 
in  the  USNM. 

Comments.  O.  c.  pansa  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  513). 


Type  Specimens  of  Recent  Mammals  •  Helaen  and  McFadden        111 


Orthogeomys  grandis  pluto  Lawrence, 

1933a 

Proc.  New  England  Zool.  Club,  13:  66,  8 

May. 

Holotype.  MCZ  29040.  Skin  and  skull.  Adult  female. 
Locality.   Honduras:  Francisco  Morazan,  north  of 
Tegucigalpa,  Cerro  Cantoral.  20  July  1932. 
Collector.    C.    F.    Underwood.    Original    number 
1100. 

Condition.  Skin  and  skull  complete. 
Type   Series.   2  paratypes;   MCZ  29038,  skin  and 
sk-ull,  juvenile  female;  MCZ  29039,  skin  and  skull 
unsexed  juvenile. 

Coninient.s.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  509). 

Genus  PAPPOGEOMYS  Mernam,  1895 

Cratogeomys  castanops  rubellus  Nelson 

and  Goldman,  1934a 

Proc.  Biol.  Soc.  Washington,  47:  147,  13 

June. 

=  Pappogeomys  castanops  rubellus 

(Nelson  and  Goldman,  1934).  See  Russell 

(1968b:  682). 

Holotype.  MCZ  20507.  Skin  and  skrdl.  Adult  male. 
Locality.   Mexico:  San  Luis  Potosi,  near  San  Luis 
Potosi,  Soledad,  6,400  ft  (1,952  m).  1  August  1923. 
Collector  W.  W.  Browai,  Jr.  Original  number  2.31. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  There  is  a  large  paratype  series  in  the 
MCZ. 

Comments.  P.  a.   nibellus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  520). 


Family  HETEROMYIDAE  Gray,  1868 

Genus  DIPODOMYS  Gray,  1841 

Dipodomys  californicus  pallidulus  Bangs, 

1899J 

Proc.  New  England  Zool.  Club,  1:  65,  31 

July. 

=  Dipodomys  californicus  californicus 

Merriam,  1890.  See  Kelt  (1988:  1). 

Holotype.  B9147.  Skin  and  skull.  Adult  female. 
Locality.  (United  States):  California,  Colusa  Coun- 
ty, Sites.  27  June  1896. 

Collector  P.  O.  Simons.  Original  number  222. 
Condition.   Skin  complete.  Skull  partial  (left  jugal 
missing). 

Type  Series.  1  paratype;  B9148,  sldn  and  skull, 
adult  female. 

Dipodops  ordii  palmeri  J.  A.  Allen,  1891 

Bull.  Amer.  Mus.  Nat.  Hist.,  3:  276,  30 

June. 

=  Dipodomys  ordii  palmeri  {y}.  A.  Allen, 

1891).  See  Grinnell  (1921:  96). 

Syntypes.  MCZ  5886:  Skin  and  skull.  Juvenile  male. 
MCZ  5887:  Skin  and  skull.  Unsexed  juvenile. 
Locality.  Mexico:  San  Luis  Potosi,  San  Luis  Potosi. 
1  May  1878. 
Collector  E.  Palmer. 

Condition.  MCZ  5886:  Sldn  complete.  Skull  partial 
(part  of  parietal,  right  occipital  condyle,  and  tym- 
panic bulla  missing).  MCZ  5887:  Skin  complete. 
Skull  partial  (left  squamosal  process  missing).  Man- 
dible disarticulated. 
Type  Series.  2  syntypes  only. 

Comments.  Allen  did  not  specify  a  holotype  in  the 
original  description,  which  he  based  on  the  two 
specimens  above.  D.  o.  palmeri  was  retained  as  a 
vahd  subspecies  by  Hall  (1981:  569). 


Genus  THOMOMYS  Wied-Neuwied,  1839       Genus  HETEROMYS  Desmarest,  1817 

Heteromys  repens  Bangs,  1902b 
Bull.  Mus.  Comp.  Zool,  39:  45,  April. 
^Heteromys  desmarestianus  repens 
Bangs,  1902.  See  Goldman  (1920:  115). 


Thomomys  umbrinus  atrodorsalis  Nelson 

and  Goldman,  1934b 

J.  Mammal.,  15:  111,  15  May. 

Holotype.  MCZ  20487.  Skin  and  skull.  Adult  male. 
Locality.   Mexico:  San  Luis  Potosi,  Alvarez,  8,000 
ft  (2,440  m).  7  November  1923. 
Collector  W.  W.  Brown,  Jr.  Original  number  338. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  There  is  a  large  series  of  paratypes  in 
the  MCZ. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  476). 


Holotype.  MCZ  10356.  Skin  and  skull.  Adult  female. 
Locality.   Panama:  Chiriqui,  Boquete  (  =  Bajo  Bo- 
quete),  4,000  ft  (1,220  m).  8  April  1901. 
Collector  W.  W.  Brown,  Jr.  Original  number  264. 
Condition.  Sldn  complete.  Skull  partial  (left  jugal 
and  process  of  squamosal  missing). 
Type  Series.  5  paratypes;  MCZ  10359,  adult  male, 
10355,  adult  male,   10358,  adult  male,  10361,  ju- 
venile female,  10360,  juvenile  female. 
Comments.   H.   d.   repens  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  597). 


112         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Genus  LIOMYS  Uernam,  1902 

Heteromys  alleni  Coues,  1881 

In  J.  A.  Allen,  Bull.  Mus.  Comp.  Zool.,  8: 

187,  March. 

=  Liomys  irroratus  alleni  (Coues,  1881). 

See  Goldman  (1911:  56). 

Holotype.  MCZ  5889.  Skin  and  skull  Male. 

Locality.  (Mexico):  San  Luis  Potosi,  Rio  Verde,  Ha- 
cienda Angostura.  26  Febniaiy  1878. 
Collector.  E.  Palmer. 

Condition.  Sldn  complete,  skull  withiji  skin. 
Type  Series.  Holotype  only. 

Comments.  L.  i.  alleni  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  590). 

Genus  PEROGNATHUSW\e6-NeuW\e6, 
1839 

Perognathus  longimembris  bangs!  Mearns, 

1898 

Bull.  Amer.  Mus.  Nat.  Hist.,  10:  300,  31 

August. 

Holotype.  B5304.  Skin  and  skull.  Adult  female. 

Locality.  (United  States):  California,  (Riverside 
County),  Colorado  Desert,  Palm  Springs,  450  ft 
(137  m).  13  April  1896. 

Collector.  E.  C.  Thurber.  Original  number  644. 
Condition.  Sldn  complete.  Skull  partial  (teeth  sep- 
arate from  skull).  Mandible  disaiUculated. 
Type  Series.  2  paratypes;  B5302,  adult  male,  skin 
and  sloill;  B5303,  adult  female,  skin  and  skull. 
Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  537). 

Family  DIPODIDAE  Fischer  de  Waldheim, 
1817 

Genus  ZAPUS  Coues,  1875 

Zapus  hudsonius  hardy!  Batchelder,  1 899 

Proc.  New  England  Zool.  Club,  1:  5,  8 

February. 

=Zapus  hudsonicus  acadlcus  (Dawson, 

1856).  See  Krutzsch  (1954:  432). 

Holotype.  MCZ  41681.  Skin  and  skull.  Adult  female. 
Locality.  (United  States):  Maine,  Hancock  County, 
Mount  Desert  Island.  24  August  1898. 
Collector  C.  F.  Batchelder.  Original  number  1597. 
Condition.  Skin  and  skull  complete. 
Tijpe  Series.  There  is  a  series  of  paratypes  in  tlie 
MCZ. 


Zapus  hudsonius  ladas  Bangs,  1 899d 
Proc.  New  England  Zool.  Club,  1:  10,  28 
February. 

Holotype.  B4169.  Skin  and  skull.  Adult  female. 
Locality.  (Canada):  Labrador,  Hamilton  Inlet,  Ri- 
goulette.  18  July  1895. 

Collector.  C.  H.  Goldthwaite.  Original  number  2. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  There  is  a  large  series  of  paratypes, 
most  of  which  are  in  the  MCZ. 
Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  843). 

Zapus  orahus  Preble,  1899 
N.  Amer.  Fauna,  15:  29,  8  August. 
=Zapus  trinotatus  orarlus  Preble,  1899. 
See  Hooper  (1944:  67). 

Holotype.  B250.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  California,  Marin  Coun- 
ty, Point  Reyes.  14  May  1893. 
Collector  C.  A.  Allen.  Original  number  618. 
Condition.  Skin  and  skull  complete. 
Type  Series.  3  paratypes,  not  in  MCZ;  mentioned 
by  locality  in  original  description. 
Comments.  Z.  t.  orarius  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  846)  and  Gannon  (1988:  1). 

Family  MURIDAE  llliger,  1811 

Subfamily  ARVICOLINAE  Gray,  1821 

Genus  CH/OA/O/V/yS  Miller,  1908 

Hypudaeus  nivlcola  Schinz,  1845 

Syst.  Verzeichniss  Saugethiere  Synopsis 

Mammalium,  2:  236. 

=  Chionomys  nivalis  (Martins,  1842).  See 

Musser  and  Carleton  (1993:  507). 

Syntype.  MCZ  1291.  Skin.  Juvenile  female. 
Locality.  Switzerland:  St.  Gotthard. 
Collector  Received  from  L.  Agassiz, 
Condition.  Sldn  complete. 

Type  Series.  Schinz  wrote  that  he  had  e.xamined  16 
specimens,  "junge  und  alte." 

Comments.  G.  M.  Allen  discussed  the  somewhat 
uncertain  tyjie  status  of  this  specimen  (1931:  263). 
The  original  label  bears  the  following:  "Hypodaeus 
[sic]  nivicola  juv.  9  sp.  nov.  du  St.  Gotthard  et  du 
Faulhorn."  Louis  Agassiz,  a  student  of  Schinz's,  de- 
posited the  specimen  in  the  MCZ. 


Type  Specimens  of  Recent  Mammals  •  Helgen  and  McFadden        113 


Genus  CLETHRIONOMYSJWes'wJs,  1850        Genus  EOTHENOMYS  Miller,  1896 


Evotomys  proteus  Bangs,  1 897g 

In  Bailey,  Proc.  Biol.  Soc.  Washington, 

11:  137,  13  May. 

=  Clethrionomys  gapperi  proteus  (Bangs, 

1897).  See  Jackson  (1938:  433). 

Holotype.  B4081.  Sldn  and  skull.  Adult  female. 

Locality.  (Canada):  Labrador,  Hamilton  Inlet.  27 
August  1895. 

Collector.  C.  H.  Goldthwaite.  Original  number  10. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  There  is  a  large  series  of  paratypes  in 
tlie  MCZ. 

Comments.  C.  g.  proteus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  783). 

Genus  DICROSTONYX  Gloger,  1841 

Dicrostonyx  chionopaes  G.  M.  Allen, 

1914b 

Proc.  New  England  Zool.  Club,  5:  62,  9 

April. 

=  Dicrostonyx  torquatus  chionopaes  G.  M. 

Allen,  1914.  See  Ognev  (1948:  507). 

Holotype.  MCZ  15263.  Skin  and  skull.  Adult  male. 
Locality.  U.S.S.R.  (  =  Russian  Federation):  eastern 
Siberia,    Nijni    Kolymsk    (Nizhnekolymsk),    near 
mouth  of  Kolyma  River.  15  October  1911. 
Collector  J.  Koren.  Original  number  257. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 
Type  Series.  Holotype  only. 

Dicrostonyx  exsuKj.  M.  Allen,  1919a 
Bull.  Mus.  Comp.  Zool.,  62:  532, 
February. 

Holotype.  MCZ  11885.  Skin  and  skull.  Adult  male. 
Locality.  (United  States):  (Alaska),  St.  Lawi-ence  Is- 
land. 24  June  1913. 

Collector  J.  Dixon.  Original  number  3267. 
Condition.  Skin  complete.  Skull  partial  (basioccip- 
ital  and  right  tympanic  bulla  missing).   Mandible 
disarticulated. 

Type  Series.  3  paratypes;  MCZ  11883,  skin  and 
skull,  female;  MCZ  11884,  skin  and  skull,  female; 
USNM  232007,  skin  and  skull,  female. 
Comments.  Considered  a  valid  species  by  Musser 
and  Carleton  (1993:  510)  but  included  in  D.  groen- 
landicus  by  Nov^^ak  (1999:  1479). 


Craseomys  aquilus  G.  M.  Allen,  1912b 

Mem.  Mus.  Comp.  Zool,  40:  216, 

August. 

=  Eotlienomys  eva  eva  (Thomas,  1911). 

See  G.  M.  Allen  (1940:  837). 

Holotype.  MCZ  7190.  Skin  and  skull.  Adult  male. 
Locality.   China:   Hupeh  (  =  Hubei),  Showlungtan. 
17  May  1907. 

Collector  W.  R.  Zappey  Original  number  10. 
Condition.  Sldn  complete.  Skull  partial  (hole  in  left 
parietal,  occiput  missing). 

Type  Series.  5  paratypes;  MCZ  7189,  7191-7194, 
7196;  all  represented  by  sldn  and  skull,  all  female. 

Microtus  (Eothenomys)  aurora  G.  M. 

Allen,  1912b 

Mem.  Mus.  Comp.  Zool.,  40:  211, 

August. 

=  Eotlienomys  melanogaster  aurora  (G.  M. 

Allen,  1912).  See  Hinton  (1923:  149). 

Holotype.  MCZ  7788.  Skin  and  skull.  Male. 

Locality.    China:   Hupeh  (  =  Hubei),  Changyangh- 

sieh.  2  Februaiy  1909. 

Collector  W.  R.  Zappey.  Original  number  372. 

Condition.  Skin  and  skull  complete. 

Type  Series.  3  paratypes;  MCZ  7185,  sldn  and  skull, 

female;    MCZ  7186,   skin   and  skull,   male;   MCZ 

7188,  skin  and  skull,  male. 

Comments.  E.  ni.  aurora  was  retained  as  a  valid 

subspecies  by  Zhang  et  al.  (1997:  229). 

Microtus  (Eotlienomys)  mucronatus  G.  M. 

Allen,  1912b 

Mem.  Mus.  Comp.  Zool.,  40:  214, 

August. 

=  Eotiienomys  melanogaster  melanogaster 

(Milne-Edwards,  1871).  See  G.  M.  Allen 

(1940:  806). 

Holotype.  MCZ  7789.  Sldn  and  skull.  Aduk  female. 
Locality.    China:    western    Szechwan    (  =  Sichuan), 
Tachiao,  12,000  ft  (3,660  m).  11  August  1908. 
Collector.  W.  R.  Zappey.  Original  number  269. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  3  paratypes;  MCZ  7790,  sldn  and  skull, 
adult  female  (exchanged  to  the  BMNH);  MCZ 
7791,  skin  and  skuU,  adult  female;  MCZ  7803,  skin 
and  skuU,  juvenile  female. 


114         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Genus  LEMMUS  Link,  1795 

Lemmus  paulus  G.  M.  Allen,  1914b 
Proc.  New  England  Zool.  Club,  5:  60,  9 
April. 

=  Lemmus  sibiricus  chrysogasterJ.  A. 
Allen,  1903.  See  Ellerman  and  Morrison- 
Scott  (1951:  656). 

Holotijpe.  MCZ  15268.  Skin  and  skull.  Adult  male. 
Locality.  (Russian  Federation):  northeastern  Sibe- 
ria, Kalascliowo,  near  mouth  of  Kolyma  River.  22 
June  1912. 

Collector.  J.  Koren.  Original  number  152. 
Condition.  Skin  complete.  Skull  partial  (left  man- 
dibular ramus  missing). 
Ti/pe  Series.  1  paratype;  MCZ  15267,  skin,  female. 

Genus  M IC ROTUS  Schrank,  1798 

Arvicola  breweri  Bah6,  1858 
Mammals,  in  Repts.  Explor.  Surveys 
Railr.  to  Pacific,  8(1):  525,  14  July. 
=  Microtus  breweri  {Bai'cd,  1858).  See 

Miller  (1896:  83). 

Syntype.    MCZ   4365.    Body   in    alcohol.    Formerly 
USNM  2833. 

Locality.  (United  States);  Massachusetts,  Nantuck- 
et County,  off  Nantucket  Island,  Muskeeget  (  =  Mu- 
skeget)  Island.  July  1856. 
Collector.  T.  M.  Brewer. 
Condition.  Alcoholic. 

Type  Series.  There  are  5  other  syntypes  (see  Poole 
and  Schantz,  1942:  271-272),  all  of  which  were  at 
one  time  in  the  USNM. 

Comments.  Poole  and  Schantz  (1942:  271)  noted 
that  USNM  2829,  a  svntype  of  breweri,  could  not 
be  found. 

Arvicola  riparia  var.  longipilis  Baird,  1858 
Mammals,  in  Repts.  Explor.  Sin-vevs 
Railr.  to  Pacific,  8(1):  524,  14  July' 
=  Microtus  pennsylvanicus  pennsylvanicus 
(Ord,  1815).  See  Bailey  (1900:  16). 

Syntype.  MCZ  5292.  Skin  and  skull. 

Locality.  (United  States):  Illinois,  (Cook  County), 
West  Northfield.  Spring  1855. 
Collector  R.  Kennicott.  Formerly  USNM  745. 
Condition.  Sldn  complete.  Skull  partial  (part  of  pa- 
rietal, both  tympanic  bullae  and  palatine  missing). 
Mandible  disarticulated. 
Tijpe  Series.  See  comments. 

Comments.  G.  M.  Allen  (1931:  260),  mentioned 
that  "although  no  individuals  are  mentioned  [in 
Baird's  description],  .  .  .  [tliis]  specimen  was  un- 
doubtedly among  those  examined  by  Baird  in  pre- 
paring his   diagnosis   and,   therefore,   is   a  cotype 


[  =  syntype]."  Baird's  specimens  were  collected  by 
Kennicott  at  West  Northfield,  Illinois,  and  by  Hoy 
at  Racine,  Wisconsin,  and  were  apparently  depos- 
ited in  the  USNM.  Lyon  and  Osgood  (1909)  and 
Poole  and  Schantz  (1942)  make  no  mention  of  oth- 
er sxii  types. 

Arvicola  rufidorsum  Baird,  1858 

Repts.  Explor.  Surveys  Railr.  to  Pacific, 

8(1):  526,  14  July. 

=  Microtus  pennsylvanicus  pennsylvanicus 

(Ord,  1815).  See  Bailey  (1900:  16). 

Holotype.  MCZ  54372.  Skin  and  skull. 

Locality.   (United  States):   Massachusetts,  (Duke's 
County),  Martha's  Vineyard,  Holmes  Hole. 
Collector  D.  J.  Wyman. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Holotype  only. 

Comments.  This  specimen  has  formerly  been 
BSNH  1949  and  USNM  901. 

Arvicola  terraenovae  Bangs,  1894a 

Proc.  Biol.  Soc.  Washington,  9:  129,  27 

July. 

=  Microtus  pennsylvanicus  terraenovae 

(Bangs,  1894).  See  Davis  (1936:  290). 

Holotype.  B1104.  Skin  and  skull.  Adult  male. 

Locality.  (Canada):  Newfoundland,  Codroy.  27  No- 
vember 1893. 

Collector  E.  Doane.  Original  number  4. 
Condition.  Sldn  and  sk-ull  complete. 
Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  M.  p.  terraenovae  was  retained  as  a  val- 
id subspecies  bv  Hall  (1981:  796). 

Microtus  chrotorrhinus  ravus  Bangs, 

1898p 

Proc.  Biol.  Soc.  Washington,  12:  188,  16 

November. 

Holotype.  B7951.  Skin  and  skull.  Adult  male. 

Locality.   (Canada):  Labrador,  Strait  of  Belle  Isle, 
Black  Bay.  15  July  1898. 
Collector.  E.  Doane.  Original  number  4. 
Condition.  Skin  and  skull  coiuplete.  Mandible  dis- 
aiticulated. 

Type  Series.  Paratype  mateiial  consists  of  Bangs' 
series  from  the  type  locahty;  B7952-7967,  all  rep- 
resented by  skin  and  skull,  10  females,  3  males,  and 
3  unsexed  individuals.  B7953  and  B7955  v^ere  ex- 
changed to  MVZ  in  1937;  B7964  was  exchanged  to 
USNM  in  1922. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  811)  and  Kirkland  and  Jannett  (1982:'  1). 


Type  Specimens  of  Recent  Mammals  •  Helaen  and  McFadden 


115 


Microtus  enixus  Bangs,  1896g 
Amer.  Nat.,  30:  1051,  1  December. 
=  Microtus  pennsylvanicus  enixus  Bangs, 
1896.  See  Davis  (1936:  290). 

Holotype.  B3973.  Skin  and  skull.  Adult  female. 
Locality.   (Canada):  Labrador,  Hamilton  Inlet.  15 
July  1895. 

Collector.  C.  H.  Goldthwaite.  Original  number  4. 
Condition.  Skin  and  skull  complete. 
Type  Series.  There  is  a  large  series  of  parat\'j3es  in 
the  MCZ. 

Comments.  M.  p.  enixus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  79.3). 

Microtus  fontigenus  Bangs,  1896d 

Proc.  Biol.  Soc.  Washington,  10:  48,  9 

March. 

=  Microtus  pennsylvanicus  fontigenus 

Bangs,  1896.  See  Miller  (1897:  14). 

Holotype.  B3837.  Skin  and  skull.  Adult  female. 

Localitij.  (Canada):  Quebec,  Lake  Edward.  28  Sep- 
tember 1895. 

Collector.  E.  A.  and  O.  Bangs.  Original  number  9. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Tijpe  Series.  7  paratypes;  B3838-B3844;  all  repre- 
sented by  sldn  and  skull,  2  females  and  5  males. 
Comments.  M.  p.  fontigenus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  793). 

Microtus  l<oreni  G.  M.  Allen,  1914b 

Proc.  New  England  Zool.  Club,  5:  64,  9 

April. 

=  Microtus  oeconomus  koreni  G.  M.  Allen, 

1914.  See  Ellerman  and  Morrison-Scott 

(1951:  706). 

Holotype.  MCZ  15213.  Skin  and  skull.  Adult  female. 
Locality.  (Russian  Federation):  northeastern  Sibe- 
ria, Nijni  Kolymsk  (  =  Nizhnekolymsk),  near  mouth 
of  Kolyma  River.  1  November  1911. 
Collector  J.  Koren.  Original  number  132. 
Condition.  Skin  complete.  Skull  partial  (left  tym- 
panic bulla  broken,  right  mandibular  ramus  miss- 
ing). 

Type  Series.  There  is  a  large  series  of  paratypes  in 
the  MCZ. 

Microtus  pennsilvanicus  [sic]  sliattucl<i 

Howe,  1901 

Proc.  Portland  Soc.  Nat.  Hist.,  2:  201,  31 

December. 

=  Microtus  pennsylvanicus  shattucki 

Howe,  1901.  See  Wyman  (1922:  162). 

Holotype.  MCZ  10011.  Sldn  and  skull.  Adult  female. 
Locality.   (United  States):  Maine,  Penobscot  Bay, 


near  Long  Island,  Tvmible  Down  Dick  Island.  10 
July  1900. 

Collector  R.  H.  Howe  and  G.  C.  Shattuck.  Original 
number  31. 

Condition.  Skin  and  skull  complete. 
Type  Series.  There  is  a  small  series  of  paratypes  in 
the  MCZ. 

Comments.  Wyman  (1922:  166)  considered  M.  p. 
shattucki  a  synonym  of  M.  p.  pennsylvanicus,  but 
Hall  (1981:  796)  retained  it  as  a  separate  subspe- 
cies. 

Microtus  pennsylvanicus  acadicus  Bangs, 
1897c 

Amer.  Nat.,  31:  239,  1  March. 

Holotype.  B2155.  Skin  and  skull.  Adult  female. 

Locality.    Canada:    Nova    Scotia,    Digbv.    22    |uly 

1894. 

Collector  O.  Bangs. 

Condition.  Sldn  and  skull  complete. 

Type  Series.  There  is  a  series  of  paratypes  in  the 

MCZ. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 

(1981:  792). 

Microtus  provectus  Bangs,  1 908 

Proc.  New  England  Zool.  Club,  4:  20,  6 

March. 

=  Microtus  pennsylvanicus  provectus 

Bangs,  1908.  See  Chamberlain  (1954: 

589). 

Holotype.  B9794.  Skin  and  skull.  Adult  female. 

Locality.  (United  States):  Rhode  Island,  Block  Is- 
land. 5  August  1899. 

Collector  O.  Bangs.  Original  number  11. 
Condition.  Sldn  complete.  Skull  partial  (tympanic 
bullae  broken).  Mandible  disarticulated  and  angu- 
lar process  chipped. 

Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  M.  p.  provectus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  796). 

Genus  MYOPUSmWer,  1910 

Myopus  thayeri  G.  M.  Allen,  1914b 

Proc.  New  England  Zool.  Club,  5:  58,  9 

April. 

^Myopus  schisticolor  thayeri  G.  M.  Allen, 

1914.  See  Ognev  (1948:  526). 

Holotype.  MCZ  15264.  Sldn  and  skull.  Adult  male. 
Locality.  (Russian  Federation):  northeastern  Sibe- 
ria,   Yakutsk,    Nijni    Kolyinsk    (  =  Nizhnekolyinsk), 
near  mouth  of  Kolyma  River  28  March  1912. 
Collector  J.  Koren.  Original  number  264. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 


116         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Comment.^.  1  paratype;  MCZ  15265,  skin  and  skull, 
male. 

Genus  ONDATRA  Link,  1795 


Fiber  obscurus  Bangs,  1894b 

Proc.  Biol.  Soc.  Washington,  9:  133,  15 

September. 

=  Ondatra  zibethicus  obscurus  (Bangs, 

1894).  See  Cameron  (1959:  85). 

Holotype.  B1155.  Skin  and  skull.  Adult  female. 

Locality.  (Canada):  Newfoundland,  Codroy.  14 
May  1894. 

Collector.  E.  Doane.  Original  number  3. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  There  is  a  small  series  of  paratypes  in 
the  MCZ. 

Comments.  O.  z.  obscurus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  827). 

Fiber  zibethicus  aquilonius  Bangs,  1899d 

Proc.  New  England  Zool.  Club,  1:  11,  28 

February. 

=  Ondatra  zibettiicus  aquilonius  (Bangs, 

1899).  See  Miller  (1912:231). 

Holotype.  B3957.  Skin  and  skull.  Adult  male. 

Locality.  (Canada):  Labrador,  Hamilton  Inlet,  Ri- 
goulette.  15  August  1895. 

Collector.  C.  H.  Goldthwaite.  Original  number  11. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  3  paratypes;  B7974,  sldn  and  skull, 
adult  female;  B7975,  sldn  and  skull,  juvenile; 
B8704,  skull,  juvenile. 

Comments.  O.  z.  aquilonius  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  825). 

Fiber  zibethicus  rivalicius  Bangs,  1895b 
Proc.  Boston  Soc.  Nat.  Hist.,  26:  541,  31 

=  Ondatra  zibethicus  rivalicius  (Bangs, 
1895).  See  Davis  and  Loweiy  (1940: 

212). 

Holotype.  B2719.  Sldn  and  skull.  Adult  male. 

Locality.  (Canada):  Louisiana,  Plaquemines  Parish, 

Burbridge.  31  January  1895. 

Collector.  F.  L.  Small.  Original  number  1556/165. 

Condition.  Sldn  and  skull  complete. 

Type  Series.   4  paratypes;   B2720,  skin  and  skull, 

adult  female;   B2721,  skin  and  skull,  adult  male; 

B2883,  skin  and  skull,  male;  B2884,  sldn  and  skull, 

male. 

Comments.  O.  z.  rivalicius  was  retained  as  a  valid 

subspecies  by  Hall  (1981:  827). 


Genus  PHENACOMYS  Mernam,  1889 

Phenacomys  celatus  crassus  Bangs, 

1900c 

Proc.  New  England  Zool.  Club,  2:  39,  20 

September. 

=  Phenacomys  ungava  crassus  Bangs, 

1900.  See  A.  B.  Howell  (1926:  27). 

Holotype.  B3959.  Skin  and  skull.  Adult  male. 

Locality.   (Canada):  Labrador,  Hamilton  Inlet,  Ri- 

goulette.  15  August  1895. 

Collector  C.  H.  Goldthwaite.  Original  number  1. 

Condition.  Sldn  and  skull  complete. 

Type  Series.  There  is  a  small  series  of  paratypes, 

most  of  which  are  no  longer  in  the  MCZ. 

Comments.  In  the  original  description,  "3946"  is  a 

misprint  for  3964. 

Genus  SYNAPTOMYS  Ba\r6,  1858 

Synaptomys  fatuus  Bangs,  1 896d 

Proc.  Biol.  Soc.  Washington,  10:  47,  9 

March. 

=  Synaptomys  cooperi  cooperi  Baird, 

1858.  See  Wetzel  (1955:8). 

Holotype.  B3857.  Skin  and  skull.  Adult  female. 
Locality.  (Canada):  Quebec,  Lake  Edward.  28  Sep- 
tember 1895. 

Collector  E.  A.  and  O.  Bangs.  Original  number  3. 
Condition.  Sldn  and  skull  complete. 
Type  Series.   8  paratypes;  B3854-B3856,  B3858- 
B3862;  all  represented  by  sldn  and  skull,  3  females 
and  5  males. 

Synaptomys  (Mictomys)  innuitus 

medioximus  Bangs,  1 900c 

Proc.  New  England  Zool.  Club,  2:  40,  20 

September. 

=  Synaptomys  borealis  medioximus 

Bangs,  1900.  See  A.  B.  Howell  (1927:  9). 

Holotype.  B8852.  Skin  and  skull.  Adult  male. 

Locality.   (Canada):  Labrador,  Strait  of  Belle  Isle, 
Lance  (  =  Lanse)  an  Loup.  15  April  1899. 
Collector  E.  Doane.  Original  number  7. 
Condition.   Sldn  complete.  Skull  slightly  damaged 
(hole  in  left  parietal). 

Type   Series.    1   paratype;    B3972,   sldn   and  skull, 
adult  male. 

Comments.  S.  b.  medioximus  was  retained  as  a  val- 
id subspecies  by  Hall  (1981:  834). 


Type  Specimens  of  Recent  Mammals  •  Helaen  and  McFadden        117 


Subfamily  CRICETINAE  Fischer  de 
Waldheim,  1817 

Genus  CANSUMYS  G.  M.  Allen,  1928 

Cansumys  canus  G.  M.  Allen,  1928 
J.  Mammal.,  9:  245,  9  August. 

Holotype.  MCZ  23779.  Skin  and  skull.  Adult  female. 
Locality.    China:    southern    Kansu    (=Xinjiang), 
Choni.  9  December  1925. 
Collector.  R.  B.  Ekvall.  Original  number  79. 
Condition.  Skin  complete.  Skull  partial  (left  jugal 
and  occiput  missing). 

Type  Series.  1  paratype;  MCZ  23780,  skin  and 
skull,  juvenile  male. 

Comments.  Type  species  o( Cansumys  G.  M.  Allen, 
1928.  Considered  a  valid  species  by  Musser  and 
Carleton  (1993:  537)  and  Nowak  (1999:  1423). 
Known  only  by  the  type  series  and  one  additional 
specimen,  FMNH  36067. 

Subfamily  CRICETOMYINAE  Roberts, 
1951 

Genus  SACCOSTOMUS  Peters,  1846 

Saccostomus  cricetulus  G.  M.  Allen  and 
Lawrence,  1936 

Bull.  Mus.  Comp.  Zool.,  79:  100,  January. 
=  Saccostomus  mearnsiHeWer ,  1910.  See 
Hubert  (1978:  51). 

Holotype.  MCZ  31475.  Skin  and  skaill.  Subadult  male. 
Locality.    Uganda:    Sabei    District,    due    north    of 
Mount  Elgon,  south  bank  of  Greek  River  (  =  Kelim 
River),  3,000  ft  (915  m).  5  December  1933. 
Collector.  A.  Loveridge. 

Condition.  Skin  complete.  Skull  partial  (right  tym- 
panic bulla  missing).  Mandible  disarticulated. 
Type   Series.    1   paratype;    MCZ  31474,   skin   and 
skull,  adult  female. 

Subfamily  DENDROMURINAE  G.  M. 
Allen,  1939 

Genus  S7E/\ rO/WVS  Peters,  1846 

Steatomys  pratensis  nyasae  Lawrence 

and  Loveridge,  1953 

Bull.  Mus.  Comp.  Zool,  110:  39,  June. 

Holotype.  MCZ  44213.  Skin  and  skull.  Adult  male. 
Locality.    Nyasaland  (  =  Malawi):   foot  of  Mulanje 
Mountain,  Likabula  River,  2,100  ft  (641  m).  29  July 
1948. 

Collector.  A.  Loveridge. 
Condition.  Sldn  and  skull  complete. 
Type    Series.    14    paratypes;    MCZ   44214-44216, 
44218-44228;  all  represented  by  skin  and  skull,  9 
males  and  5  females. 


Comments.  Retained  as  a  valid  subspecies  by  An- 
sell  (1978:  77). 

Subfamily  GERBILLINAE  Gray,  1825 

Genus  TATERA  Lataste,  1882 

Tatera  flavipes  G.  M.  Allen,  1914d 
Bull.  Mus.  Comp.  Zool.,  5S:  331,  July. 
=  Tatera  valida  /cemp/ Wroughton,  1906. 
See  Bates  (1988:  277). 

Holotype.  MCZ  14491.  Skin  and  skull.  Adult  female. 
Locality.  Sudan:  Blue  Nile,  north  of  (Er)  Roseires, 
Aradeiba.  22  Januaiy  1913. 

Collector  G.  M.  Allen  and  J.  C.  PhilHps.  Original 
number  69. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 
Type  Series.  Holotype  only. 

Tatera  sororG.  M.  Allen,  191 4d 
Bull.  Mus.  Comp.  Zool,  58:  333,  July 
=  Tatera  valida  /cemp/ Wroughton,  1906. 
See  Bates  (1988:  277). 

Holotype.  MCZ  14492.  Skin  and  skull.  Adult  female. 
Locality.  Sudan:  Blue  Nile,  Fazogli.  16  January 
1913. 

Collector  CM.  Allen  and  J.  C.  Phillips.  Original 
number  53. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type   Series.    1   paratype;    MCZ    14493,   skin   and 
skull,  juvenile  male. 

Genus  TATERILLUS  Thomas,  1910 

Taterillus  melanops  G.  M.  Allen,  1912a 
Bull.  Mus.  Comp.  Zool.,  54:  446,  April. 
=  Taterillus  harringtoni  (Thomas,  1906). 
See  Musser  and  Carleton  (1993:  563). 

Holotype.  MCZ  8132.  Skin  and  skull.  Male. 

Locality.  British  East  Africa  (  =  Kenya):  arid  plains 
by  the  Meru  River.  11  August  1909. 
Collector  G.  M.  Allen.  Original  number  108. 
Condition.  Skin  and  skull  complete. 
Tijpe  Series.  Holotype  only. 

Subfamily  MURINAE  llliger,  1811 

Genus  APODEMUS  Kaup,  1829 

Apodemus  mystacinus  euxinus  G.  M. 

Allen,  1915b 

Bull.  Mus.  Comp.  Zool,  59:  11, 

Februaiy. 

= Apodemus  mystacinus  mystacinus 

(Danford  and  Alston,  1877).  See  Corbet 

(1978:  133). 

Holotype.  MCZ  14887.  Skin  and  skull.  Male. 

Locality.   Asia  Minor  (=Turkey):   (Trabzon),  near 


118         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Trebizond  (=Trabzon),   Scalita,   1,000  m.  25  No- 
vember 1905. 

Collector.  A.  Robert.  Original  number  2189. 
Condition.  Skin  complete.  Skull  partial  (right  and 
left  jugal  missing,  right  tympanic  bulla  separated 
but   present,    right   and   left   squamosal    missing). 
Mandible  disarticulated. 
Tijpe  Series.  HolotN-pe  only. 

Genus  DASYMYS  Peters,  1875 

Dasymys  incomtus  alleni  Lawrence  and 

Loveridge,  1953 

Bull.  Mus.  Comp.  ZooL,  110:  53,  June. 

Holotijpe.  MCZ  26322.  Skin  and  skull.  Adult  male. 
Locality.      Tanganyika     Territoiy     (=Tanzania): 
(Mbeya),  Ilolo,  near  Rungwe  Mountains,  4,600  ft 
(1,403  m).  31  March  1930. 
Collector.  A.  Loveridge. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated and  right  coronoid  process  chipped. 
Type  Series.   11  paratypes,  all  represented  by  sldn 
and  skull,  in  the  MCZ. 

Comments.  Retained  as  a  valid  subspecies  by  An- 
sell  (197S:  S3). 

Genus  GRAMMOMYS  Jhomas,  1915 

Thamnomys  ochraceus  G.  M.  Allen, 

1912a 

Bull.  Mus.  Comp.  ZooL,  54:  442,  April 

1912. 

=  Grammomys  macmillani  (Wroughton, 

1907).  See  Musser  and  Carleton  (1993: 

594). 

Holotype.  MCZ  8126.  Skin  and  skull.  Adult  male. 
Locality.  British  East  Africa  (  =  Kenya):  Meru  Riv- 
er,   near  junction   with    northern    Guaso    Nyiro 
(  =  Ewaso  Ngiro).  8  August  1909. 
Collector.  G.  M.  Allen.  Original  number  103. 
Condition.  Skin  and  skull  complete. 
Type  Series.  Holotype  only. 

Genus  HyLO/WySCL/S  Thomas,  1926 

Hylomyscus  alleni  simus  G.  M.  Allen  and 

Coolidge,  1930 

7/7  Strong,  The  African  Republic  of 

Liberia  and  the  Belgian  Congo,  2:  599, 

October. 

=  Hylomyscus  alleni  (Waterhouse,  1838). 

See  Heim  de  Balsac  and  Aellen  (1965: 

722). 

Holotype.  MCZ  24028.  Skin  and  skull.  Adult  male. 
Locality.  Liberia:  Merikay.  13  September  1926. 


Collector  G.  M.  Allen  and  H.  J.  Coolidge,  Jr.  Orig- 
inal number  97. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Genus  MELO/W/S  Thomas,  1922 

Melomys  levipes  stevensi  Rummler,  1935 

Z.  Saugetierkunde,  10:  109,  31 

December. 

=  Melomys  mollis  Thomas,  1913.  See 

Flanneiy  (1990:  226). 

Holotype.  MCZ  29890.  Skin  and  skull.  Adult  male. 
Locality.  Papua  New  Guinea:  Morobe,  Mount  Mis- 
im  (  =  Missim),  6,700  ft  (2,044  m).  17  April  1933. 
Collector  H.  Stevens. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  11  paratypes;  MCZ  29889,  29892- 
29899,  29901;  all  represented  by  sldn  and  skull,  4 
females  and  7  males. 

Melomys  moncktoni  alleni  Rummler,  1935 

Z.  Saugetierkunde,  10:  112,  31 

December. 

=  Melomys  rubex  alleni  Rummler,  1935. 

See  Tate  (1951:  299). 

Holotype.  MCZ  29902.  Skin  and  skull.  Adult  female. 
Localiti/.  Papua  New  Guinea:  Morobe,  Mount  Mis- 
im  (  =  Missim),  6,700  ft  (2,044  m).  17  April  1933. 
Collector  H.  Stevens. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  1  paratvpe;  MCZ  29900,  skin  and 
skull,  adult  male. 

Comments.  Flannery  (1995a:  308—309)  did  not  list 
subspecies  of  M.  nibex,  noting,  "It  is  a  veiy  variable 
species,  and  a  taxonomic  revision  is  needed  in  or- 
der to  clarify  the  taxonomic  status  of  many  popu- 
lations." 

Genus  MUS  Linnaeus,  1758 

Leggada  bufo  ablutus  G.  M.  Allen  and 

Loveridge,  1942 

Bull.  Mus.  Comp.  ZooL,  89:  199, 

Februaiy. 

=  Mus  bufo  (Thomas,  1906).  See  Musser 

and  Carleton  (1993:  622). 

Holotype.  MCZ  40745.  Skin  and  skull.  Adult  male. 
Locality.   Belgian  Congo  (  =  Democratic  Republic 
of  Congo):  Kivu,  Lake  Kivu,  Idj\\d  Island,  Upper 
Mulinga,  6,500  ft  (1,983  m).  24  February  1939. 
Collector  A.  Loveridge. 


Type  Specimens  of  Recent  Mammals  •  Helgen  and  McFadden        119 


Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Tijpe  Series.  2  paratypes;  MCZ  40746,  skin  and 
skull  (no  longer  in  MCZ);  MCZ  40747,  skin  and 
skull,  female. 

Leggada  gerbillus  G.  M.  Allen  and 

Loveridge,  1933 

Bull.  Mus.  Comp.  ZooL,  75:  112, 

Febriiaiy. 

=  Mus  tenellus  (Thomas,  1903).  See 

Musser  and  Carleton  (1993:  629). 

Holotype.  MCZ  26586.  Skin  and  skull.  Adult  male. 
Locality.    Tanganyika  Territoiy  (=  Tanzania):    Do- 
doma,  Ugogo,  3,700  ft  (1,129  m).  23  December 
1929. 

Collector.  A.  Loveridge. 

Condition.  Sldn  complete.  Skull  partial  (right  jugal 
missing).  Mandible  disarticulated  and  angular  pro- 
cess of  right  ramus  chipped. 
Type  Series.  Holotype  only. 

Mus  bactrianus  tantillus  G.  M.  Allen,  1927 
Amer.  Mus.  Novitates  270:  9,  31  May. 
=  Mus  musculus  tantillus  G.  M.  Allen, 
1927.  See  Marshall  (1977:  214). 

Holotype.  MCZ  23476.  Skin  and  skull.  Adult  female. 
Locality.  China:  Sze-chuan  (  =  Sichuan),  Wanhsien 
(=Wanxian).  14  November  1921. 
Collector  W.  W.  Granger. 

Condition.  Skin  complete.  Skull  partial  (6  frag- 
ments). Mandible  disarticulated. 
Type  Series.  There  is  a  series  of  paratypes  in  the 
AMNH,  and  1  paratype  in  the  MCZ;  MCZ  23475 
(formerly  AMNH  56403),  skin  and  skull,  male. 
Comments.  This  specimen  was  formerly  AMNH 
56416.  The  original  description  lists  AMNH  56413 
as  the  holotype,  but  this  specimen  is  a  juvenile,  not 
an  adult  female,  as  Allen  stated.  The  collection  date 
and  measurements  of  the  holotype  in  the  original 
description  fit  MCZ  23476  rather  than  any  other 
specimens  in  the  type  series  at  the  AMNH,  which 
are  all  immatures  (Lawrence  1993:  136). 

Genus  /W/O/W/S  Thomas,  1915 

Praomys  fumatus  oweni  Setzer,  1 956 

Proc.  U.S.  Nat.  Mus.,  106:  525,  28 

November. 

=  Myomys  fumatus  (Peters,  1878).  See 

Musser  and  Carleton  (1993:  631). 

Holotype.  MCZ  45883.  Skin  and  skull.  Adult  male. 
Locality.  Anglo-Egyptian  Sudan  (  =  Sudan):  Equa- 
toria  Province,  Torit  District,  Murukurun,  50  miles 
(80.5  km)  east  of  Torit,  2,000  ft  (610  m).  9  May 
1950. 


Collector  J.  S.  Owen.  Original  number  1030. 

Condition.  Skin  and  skull  complete. 

Type  Series.  There  is  a  series  of  paratypes  in  the 

MCZ. 

Genus  A//l//\/EA/rE/?  Marshall,  1976 

Epimys  zappeyi  G.  M.  Allen,  1912b 

Mem.  Mus.  Comp.  ZooL,  40:  225, 

August. 

^Niviventer  confucianus  (Milne-Edwards, 

1871).  See  Musser  and  Carleton  (1993: 

633). 

Holotype.  MCZ  7607.  Skin  and  skull.  Adult  male. 
Locality.    China:    western    Szechwan    (  =  Sichuan), 
Washan  Mountains  (=Wu  Shan),  9,000  ft  (2,745 
m).  26  October  1908. 

Collector  W.  R.  Zappey.  Original  number  305. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 
Type  Series.  Holotype  only. 

Genus  OEA/OM/S  Thomas,  1904 

Oenomys  hypoxanthus  talangae  Setzer, 

1956 

Proc.  U.S.  Nat.  Mus.,  106:  505,  28 

November. 

Holotype.  MCZ  45315.  Skin  and  skull.  Adult  female. 
Locality.  Anglo-  Egyptian  Sudan  (  =  Sudan):  Equa- 
toria  Province,  Imatong  Mountains,  Talanga  Foi- 
est,  3,000  ft  (915  m).  10  July  1950. 
Collector  J.  S.  Owen.  Original  number  1338. 
Condition.  Skin  and  skull  complete. 
Type   Series.    1   parats-pe;    MCZ  45284,   skin   and 
skuU,  male. 


Genus  PR/A 0/WVS  Thomas,  1915 


Allen 


Praomys  tullbergi  melanotus  G. 

and  Loveridge,  1933 

Bull.  Mus.  Comp.  ZooL,  75:  106, 

February. 

=  Praomys  delectorum  (Thomas,  1910). 

See  Musser  and  Carleton  (1993:  642). 

Holotype.  MCZ  26287.  Skin  and  skull.  Adult  male. 
Locality.  Tanganyika  Territoiy  (=Tanzania):  north- 
west end  of  Lake  Nyasa,  Poroto  Mountains,  Nyam- 
wanga,  6,400  ft  (1,952  m).  21  March  1930. 
Collector.  A.  Loveridge. 

Condition.  Skin  complete.  Skull  partial  (left  tym- 
panic bulla  missing).  Mandible  disarticulated. 
Type  Series.  23  paratypes;  MCZ  26259,  26285, 
26286,  26288-26293,  26295-26297,  26387-26394, 
26411,  26497,  26498;  all  represented  by  skin  and 
skull. 


120         Bulletin  Museum  of  Comparative  Zoologi/,  Vol.  157,  No.  2 


Comments.  Van  der  Straeten  and  Dieterlen  (1987: 
9)  and  Van  der  Straeten  and  Dudu  (1990:  81)  treat- 
ed melanotus  as  a  species  within  the  Praomijs  de- 
lectorum  species  complex. 

Genus  PSEUDOHYDROMYS  Rummler, 
1934 

Pseudohydromys  murinus  Rummler,  1 934 
Z.  Saugetierkunde,  9:  48,  30  December. 

Holotype.  MCZ  29904.  Sldn  and  skull.  Adult  male. 
Locality.  Papua  New  Guinea,  Morobe,  Mount  Mis- 
im  (  =  Missim),  7,000  ft  (2,135  m).  8  March  1933. 
Collector.  H.  Stevens. 

Condition.   Sldn  complete,  with  small  bald  spots. 
Skull  complete.  Mandible  disarticulated. 
Type  Series.  Holotype  only. 

Comments.  Type  species  of  Pseudohydromys  Rii- 
mmler,  1934.  Considered  a  valid  species  by  Flan- 
nery  (1995a:  255),  Musser  and  Carleton  (1993: 
644),  and  Nowak  (1999:  1612). 

Genus  STENOMYS  Ihomas,  1910 

Stenomys  niobe  stevensi  Rummler,  1935 

Z.  Saugetierkunde,  10:  117,  31 

December. 

=  Stenomys  niobe  niobe  (Thomas,  1906). 

See  Flamieiy  (1995a:  339). 

Holotype.  MCZ  29915.  Skin  and  skull.  Adult  male. 
Locality.  Papua  New  Guinea:  Morobe,  Mount  Mis- 
im  (  =  Missim),  7,000  ft  (2,135  m).  16  Januaiy  1933. 
Collector.  H.  Stevens.  Original  number  8. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Holotype  only. 

Comments.  Taylor  et  al.  (1982:  193)  first  synony- 
mized  .stevensi  with  niobe,  under  the  genus  Rattus. 

Stenomys  verecundus  mollis  Rummler, 

1935 

Z.  Saugetierkunde,  10:  116,  31 

December. 

Holotype.  MCZ  29905.  Skin  and  skull.  Adult  female. 
Locality.  Papua  New  Guinea:  Morobe,  Mount  Mis- 
im  (  =  Missim),  5,850  ft  (1,784  m).  14  April  1933. 
Collector.  H.  Stevens. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Holotype  only. 

Comments.  Retained  as  a  valid  subspecies  by  Flan- 
neiy  (1995a:  346). 


Genus  THAMNOMYS  Thomas,  1 907 

Thamnomys  venustus  kivuensis  G.  M. 

Allen  and  Loverldge,  1942 

Bull  Mus.  Comp.  Zool,  89:  192, 

Februaiy. 

Holotype.  MCZ  39151.  Skin  and  sk-ull.  Adult  female. 
Locality.  Belgian  Congo  (  =  Democratic  Republic 
of  Congo):  (Sud-Kivu),  Lake  Kivu,  Idjwi  Island, 
Upper  Mulinga,  6,500  ft  (1,983  m).  2  March  1939. 
Collector.  A.  Loveridge. 

Condition.  Skin  complete.  Skull  partial  (separated 
between  nasal  and  frontal).  Right  and  left  angular 
processes  of  mandible  chipped. 
Tiipe  Series.  Holotype  only. 

Subfamily  NESOMYINAE  Forsyth  Major, 
1897 

Genus  NESOMYS  Peters,  1870 

Nesomys  lambertoni  G.  Grandidier,  1930c 
Bull.  Acad.  Malgache,  n.  sen,  11:  95  (for 
1928). 

Holotype.  MCZ  45941.  Skull  and  postcranial  skele- 
ton. Adult  inale. 

Locality.  Malagasy  Republic  (  =  Madagascar):  west 
coast,  vicinity  of  Maintirano. 

Collector  No  collection  data  available.  Received  by 
G.  Grandidier  from  Academic  Malgache,  1928. 
Condition.  Skull  partial  (left  zygomatic  arch  miss- 
ing, right  jugal  missing,  several  small  holes).  Post- 
cranial  skeleton  complete  (small  foot  bones  pre- 
svHTiably  preserved  in  skin). 

Type  Series.  2  paratopes,  MCZ  45933  and  45934, 
poorly  prepared  skins  (■with  partial  skulls  within), 
both  male. 

Comments.  The  skin  belonging  to  the  holotype  is 
in  the  MNHN,  Paris  (M.  Carleton,  personal  com- 
munication). Considered  a  valid  species  by  Nowak 
(1999:  1434). 

Subfamily  OTOMYINAE  Thomas,  1897 

Genus  OTOMYS  F.  Cuvier,  1824 

Otomys  anchietae  lacustris  G.  M.  Allen 

and  Loveridge,  1933 

Bull.  Mus.  Comp.  Zool.,  75:  120, 

Februaiy. 

=  Otomys  lacustris  G.  M.  Allen  and 

Loveridge,  1933.  See  Dieterlen  and  Van 

der  Straeten  (1992:  385). 

Holotype.  MCZ  26358.  Skin  and  sk-ull.  Adult  female. 
Locality.  Tanganyika  Territoiy  (=Tanzania):  north 
end  of  Lake  Nyasa,  Ukinga  Mountains,  Madehani, 
7,000  ft  (2,135  m).  21  Februaiy  1930. 


Type  Specimens  of  Recent  Mammals  •  Helgen  and  McFadden        121 


Collector.  A.  Loveridge. 

Condition.  Skin  complete.  Skiill  partial  (left  tym- 
panic bulla  damaged).  Mandible  disarticulated  and 
left  coronoid  process  chipped. 

Type  Series.  17  paratypes;  MCZ  26326,  26344- 
26351,  26353-26357,  26359,  26654,  26658;  all  rep- 
resented by  skin  and  skull. 

Comments.  Included  in  O.  anchietae  by  Musser 
and  Carleton  (1993:  680)  but  treated  as  a  full  spe- 
cies by  Nowak  (1999:  1439). 

Otomys  barbouri  Lawrence  and  Loveridge, 

1953 

BulL  Mus.  Comp.  ZooL,  110:  63,  June. 

Holotijpe.  MCZ  31369.  Skin  and  skull.  Adult  male. 
Locality.     Uganda:     Mount     Elgon,     Kaburomi, 
1°14'N,  34°31'E,  10,500  ft  (3,203  m).  28  December 
1933. 

Collector.  A.  Loveridge. 
Condition.  Skin  and  skull  complete. 
Type   Series.    9   paratypes;    MCZ   31371-31372, 
31376,  31421-31425,  31438  (4  males,  4  females, 
and  1  unsexed  individual). 

Comments.  Considered  a  valid  species  by  Dieter- 
len  and  Van  der  Straeten  (1992:  385)  and  Nowak 
(1999:  1439)  but  included  in  O.  anchietae  by  Mus- 
ser and  Carleton  (1993:  680). 

Otomys  uzungwensis  Lawrence  and 

Loveridge,  1953 

Bull  Mus.  Comp.  ZooL,  110:  61,  June. 

=  Otomys  typus  uzungwensis  Lawrence 

and  Loveridge,  1953.  See  Hanney  (1965: 

626). 

Holotype.  MCZ  26645.  Sldn  and  skull.  Adult  female. 
Locality.  Tanganyika  Territory  (=Tanzania):  Iringa 
District,    Uzungwa    Mountains,    Dadaga,   6,000   ft 
(1,830  m).  31  December  1929. 
Collector.  A.  Loveridge. 
Condition.  Skin  and  skull  complete. 
Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  O.  t.  uzungwensis  was  retained  as  a  val- 
id subspecies  bv  Ansell  (1978:  78). 

Subfamily  SIGMODONTINAE  Wagner, 
1843 

Genus  ISTHMOMYS  Hooper  and  IVIusser, 
1964 

Megadontomys  flavidus  Bangs,  1902b 
Bull  Mus.  Comp.  ZooL,  39:  27,  ApriL 
=  lsthmomys  flavidus  (Bangs,  1902).  See 
Hooper  and  Musser  (1964:  12). 

Holotype.  MCZ  10331.  Skin  and  skull.  Adult  male. 
Locality.   Panama:  Chiriqui,  Boquete  (  =  Bajo  Bo- 
quete),  4,000  ft  (1,220  m).  12  April  1901. 


Collector  W.  W.  Brown,  Jr.  Original  number  28. 

Condition.   Skin  complete,  widi  small  bald  spots. 

Skull  complete.  Mandible  disarticulated. 

Type  Series.  There  is  a  large  series  of  paratypes  in 

the  MCZ. 

Comments.  Type  species  oihthmomys  Hooper  and 

Musser,   1964.  I.  flavidus  was  considered  a  valid 

species  by  Musser  and  Carleton  (1993:  706)  and 

Nowak  (1999:  1357). 

Genus  NECTOMYS  Peters,  1861 

Nectomys  squamipes  amazonicus 
Hershkovitz,  1944 

Misc.  Pub.,  Mus.  ZooL,  Univ.  Mich.,  58: 
47,  4  Januaiy. 

Holotype.  MCZ  30820.  Skin  and  skull.  Male. 

Locality.  Brazil:  (Para),  Rio  Tapajos,  Tauary  (=Tau- 
ari).  23  January  1934. 

Collector  A.  M.  Olalla.  Original  number  7312. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Hershkovitz  examined  28  specimens, 
in  the  MCZ,  AMNH,  and  FMNH. 
Comments.  Retained  as  a  valid  subspecies  by  Er- 
nest (1986:  1). 

Genus  NEOTOMA  Say  and  Ord,  1825 

Neotoma  abbreviata  Goldman,  1909 

Proc.  BioL  Soc.  Washington,  22:  140,  25 

June. 

=  Neotoma  lepida  abbreviata  Goldman, 

1909.  See  Burt  (1932:  182). 

Holotype.  MCZ  12260.  Skin  and  skull.  Adult  male. 
Locality.  (Mexico):  Lower  (  =  Baja)  California,  San 
Francisco  Island.  22  February  1909. 
Collector.  W.  W.  Brown,  Jn  Original  number  22. 
Condition.  Skin  and  skull  complete. 
Type    Series.    9   paratypes;    MCZ    12256-12259, 
12261—12265;  all  represented  by  skin  and  skull,  5 
females  and  4  males.   MCZ  12265  is  now  in  the 
USNM. 

Comments.  N.  I.  abbreviata  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  755). 

Neotoma  bella  Bangs,  1899] 

Proc.  New  England  ZooL  Club,  1:  66,  31 

July. 

=  Neotoma  lepida  lepida  Thomas,  1893. 

See  Goldman  (1932:  62). 

Holotype.  B5308.  Skin  and  skull.  Adult  male. 

Locality.    (United  States):   California,   Riverside 
County,  Palm  Springs.  12  April  1896. 
Collector.  E.  C.  Thurber  Original  number  623. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated and  left  airgular  pr'ocess  chipped. 


122         Bulletin  Museum  of  Comparative  Zoologi/,  Vol.  157,  No.  2 


Type  Series.  Holotype  only. 

Neotoma  distincta  Bangs,  1903a 

Proc.  Biol.  Soc.  Washington,  16:  89,  25 

June. 

=  Neotoma  mexicana  distincta  Bangs, 

1903.  See  Hall  (1955:  329). 

Holotype.  B9819.  Skin  and  skull.  Adult  male. 

Locality.  Mexico:  Vera  Cruz,  near  Jalapa,  Texolo.  8 

March  1899. 

Collector.  S.  N.  Rhoads.  Original  number  295. 

Condition.  Skin  and  skull  complete. 

Type  Series.   4  paratypes;  B9818,  sldn  and  skull, 

male,  B9820,  skin  and  skull,  female;  B9821,  skin 

and  skull,  male  (exchanged  to  USNM);  B9822,  skin 

and  skull,  male. 

Comments.  N.  m.  distincta  was  retained  as  a  valid 

subspecies  by  Hall  (1981:  761)  and  Cornely  and 

Baker  (1986:  1). 

Neotoma  floridana  rubida  Bangs,  1 898b 
Proc.  Boston  Soc.  Nat.  Hist.,  28:  185,  15 
March. 

Holotype.  B2872.  Skin  and  skull.  Adult  male. 

Locality.    (United  States):   Louisiana,  Terrebonne 
Parish,  Gibson.  4  April  1895. 
Collector  F.  L.  Small.  Original  number  1751. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  2  specimens  are  mentioned  by  num- 
ber in  the  original  description:  B2871,  sldn  and 
skull,  adult  male,  and  B2873,  skin  and  skull,  adult 
female. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  749). 

Genus  NYCTOMYS  6e  Saussure,  1860 

Nyctomys  nitellinus  Bangs,  1 902b 
Bull.  Mus.  Comp.  ZooL,  39:  30,  April. 
=  Nyctomys  sumichirasti  nitellinus  Bangs, 
1902.  See  Goldman  (1916:  155). 

Holotype.  MCZ  10249.  Skin  and  skull.  Adult  female. 
Locality.   Panama:  Chiriqui,  Boquete  (  =  Bajo  Bo- 
quete),  4,000  ft  (1,220  m).  8  Februaiy  1901. 
Collector.  W.  W.  Brown,  Jr.  Original  number  119. 
Condition.  Skin  and  skull  complete. 
Type   Series.   5  paratypes;   MCZ   10245,  skin  and 
skull,  adult  male;  MCZ  10246,  skin  and  skull,  adult 
male;  MCZ   10247,  adult  female,  skin  and  skull; 
MCZ   10248,  skin  and  skull,  adult  female;   MCZ 
10250,  sldn  and  skull,  juvenile  female. 
Comments.  N.  s.  nitellinus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  630). 


Genus  OECOMYS  Thomas,  1906 

Oecomys  trabeatus  G.  M.  Allen  and 

Barbour,  1923 

Bull.  Mus.  Comp.  ZooL,  65:  262, 

Februaiy. 

=  Oecomys  bicolor  trabeatus  G.  M.  Allen 

and  Barbour,  1923.  See  comments. 

Holotype.  MCZ  19837.  Skin  and  skull.  Male. 
Locality.  Panama:  Rio  Jesusito.  10  April  1922. 
Collector.  T.  Barbour  and  W.  S.  Brooks.  Original 
number  2027. 

Condition.   Skin  complete.  Skull  partial  (right  zy- 
gomatic arch  broken).  Mandible  disarticulated. 
Type  Sei-ies.  Holotype  only. 

Comments.  Following  Hershkovitz  (1960:  533),  O. 
b.  trabeatus  was  retained  as  a  valid  subspecies  by 
Hall  (1981:  619)  under  the  genus  Oryzomys.  The 
use  of  Oecomys  as  a  generic  name  follows  Musser 
and  Carleton  (1993:  715). 

Oryzomys  flavicans  illectus  Bangs,  1898k 

Proc.  Biol.  Soc.  Washington,  12:  164,  10 

August. 

=  Oecomys  flavicans  illectus  (Bangs, 

1898k).  See  comments. 

Holotype.  B8101.  Sldn  and  skull.  Adult  female. 
Locality.    Colombia:    Magdalena,    Santa    Marta 
Mountains,  Pueblo  Viejo,  8,000  ft  (2,440  m).  24 
March  1898. 

Collector  W.  W.  Bi^own,  Jr.  Original  number  89. 
Condition.  Skin  complete.  Skull  partial  (right  jugal 
missing). 

Tt/pe  Series.  There  is  a  small  series  of  paratypes  in 
the  MCZ. 

Comments.  The  use  of  Oecomys  as  a  generic  name 
follows  Musser  and  Carleton  (1993:  715). 

Genus  OLIGORYZOMYS  Bangs,  1 900 

Oryzomys  navus  Bangs,  1899a 

Proc.  Biol.  Soc.  Washington,  13:  9,  31 

januaiy. 

=  Oligoryzomys  fulvescens  (Saussure, 

1860).  See  Carleton  and  Musser  (1989: 

70). 

Holotype.  B8107.  Skin  and  skull.  Adult  male. 

Locality.  Colombia:  Magdalena,  Santa  Marta 
Mountains,  Pueblo  Viejo,  8,000  ft  (2,440  m).  26 
March  1898. 

Collector  W.  W.  Browni,  Jr.  Original  number  154. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  9  paratypes;  B8223-B8231;  all  repre- 
sented by  sldn  and  skull,  except  B8228  (skin  only); 
2  females  and  7  males.  B8224  is  now  in  the  USNM, 


Type  Specimens  of  Recent  Mammals  *  Helpen  and  McFadden 


123 


and  B8223,  B8226,  B8230,  and  B8231  are  now  in 
the  AMNH. 

Coininents.  Oryzoini/s  navus  is  the  t)'pe  species  of 
Oligoryzomijs  Bangs,  1899. 

Oryzomys  (Oligoryzomys)  vegetus  Bangs, 

1902b 

Bull.  Mus.  Comp.  ZooL,  39:  35,  April. 

=  Oligoryzomys  vegetus  (Bangs,  1902). 

See  Musser  and  Carleton  (1993:  718). 

Holotype.  MCZ  10298.  Skin  and  skull.  Adult  female. 
Locality.    Panama:   Chiriqui,  Volcan   de   Chiriqui, 
Boquete,  4,000  ft  (1,220  m).  16  April  1901. 
Collector.  W.  W.  Brown,  Jr.  Original  number  304. 
Condition.  Skin  complete.  Skull  partial  (left  jugal 
missing).  Mandible  disarticulated. 
Type   Series.    12  paratypes;   MCZ   10295   (now  in 
USNM),  10297,  10300-10306,  10308-10310,  2  fe- 
males and  10  males,  all  represented  by  skin  and 
skull  except  for  10308  (skin  only). 
Comments.  Considered  a  valid  species  by  Musser 
and  Carleton  (1993:  718)  and  Nowak  (1999:  1368). 
Carleton  and  Musser  (1995)  provided  distribution- 
al information  and  support  of  specific  status  for  O. 
vegetus. 

Genus  OR VZOM VS  Baird,  1858 

Oryzomys  devius  Bangs,  1902b 

Bull.  Mus.  Comp.  ZooL,  39:  34,  April. 

Holotype.  MCZ  10324.  Skin  and  skull.  Adult  female. 
Locality.  Panama:  Chiriqui,  Volcan  de  Chiriqui, 
Boquete  (  =  Bajo  Boquete),  5,000  ft  (1,525  m).  29 
January  1901. 

Collector.  W.  W.  Brown,  jr.  Original  number  73. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  3  paraty^aes;  MCZ  10325,  skin  and 
skull,  adult  female;  MCZ  10326,  skin  and  skull, 
adult  female;  MCZ  10340,  sldn  and  skull,  adult 
male. 

Comments.  Considered  a  valid  species  by  Musser 
and  Carleton  (1993:  722)  and  Nowak  (1999:  1366). 

Oryzomys  palustris  coloratus  Bangs, 

1898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  189,  15 

March. 

=  Oryzomys  palustris  natator  Chapman, 

1893.  See  Wolfe  (1982:  1). 

Holotype.  B4470.  Skin  and  skull.  Adult  male. 

Localitij.  (United  States):  Florida,  Monroe  County, 

Cape  Sable.  17  April  1895. 

Collector.  C.  L.  Brownell.  Original  number  148. 

Condition.  Sldn  and  skull  complete. 

Type  Series.  There  is  a  series  of  paratypes  in  the 

MCZ. 


Oryzomys  rostratus  carrorum  Lawrence, 

1947b 

Proc.  New  England  ZooL  Club,  24:  101, 

29  May. 

Holotype.    MCZ  41280.   Skin,  skull,  and  postcranial 
skeleton.  Adult  male. 

Locality.  Mexico:  Tamaulipas,  Rio  Soto  la  Marina, 
Rancho  Santa  Ana,  8  miles  (12.9  km)  southwest  of 
Padilla.  21  December  1941. 

Collector.  G.  H.  Pournelle,  Third  University  of 
Florida  Mexican  Expedition.  Original  number  196. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. Postcranial  skeleton  partial  (right  fore- 
limb,  hindlimbs,  pelvis,  ribs,  vertebral  column). 
Type  Series.  2  paratypes;  MCZ  41281,  skin,  skull, 
and  postcranial  skeleton,  female;  MCZ  41282,  sldn, 
skull,  and  postcranial  skeleton,  male. 
Comments.  Retained  as  Oryzomys  melanotis  car- 
rorum by  Hall  (1981:  614).  Musser  and  Carleton 
(1993:  724)  regarded  O.  rostratus  as  distinct  from 
O.  melanotis. 

Genus  PEROMYSCUS  G\oger,  1841 

Hesperomys  gossypinus  LeConte,  1853 

Proc.  Acad.  Nat.  Sci.  Philadelphia,  6: 

411,  25  October. 

=  Peromyscus  gossypinus  gossypinus 

(LeConte,  1853).  See  Rhoads  (1896b: 

189). 

Lectotype.  MCZ  5275.  Skin  and  skull.  Adult  male. 
Locality.  (United  States):  Georgia,  Liberty  County, 
Riceboro,  probably  on  the  LeConte  Plantation  near 
Riceboro.  13  September  1847. 
Collector  J.  E.  LeConte.  Formerly  USNM  546. 
Condition.  Skin  complete.  Skull  partial  (basioccip- 
ital  and  tympanic  bullae  broken).  Mandible  disar- 
ticulated. 

Type  Series.  Poole  and  Schantz  (1942:  320)  listed 
potential  "cotypes,"  meaning  syntypes.  Osgood 
(1909:  136)  mentioned  that  a  specimen,  number 
752  in  the  collection  of  the  Academy  of  Natural 
Sciences  in  Philadelphia,  possibly  possessed  type 
status,  but  this  specimen  was  not  mentioned  by 
Koopman  (1976). 

Comments.  In  the  original  description  of  gossypi- 
nus, measurements  were  given  for  a  single  speci- 
men only;  however,  in  addition  to  this  specimen, 
LeConte  made  reference  to  the  coloration  of 
"younger  indixdduals."  The  measureinents  given 
correspond  to  MCZ  5275  rather  than  to  any  other 
of  LeConte "s  specimens  still  in  the  USNM  (see  G. 
M.  Allen,  1931:  262;  Poole  and  Schantz,  1942: 
320).  In  addition,  MCZ  5275  was  the  only  speci- 
men of  Peromyscus  gossypinus  from  Georgia  men- 
tioned in  Baird's  Mammals  of  North  America 
(1858:  469).  Because  LeConte  did  not  specify  a  ho- 
lotype, MCZ  5275,  which  seems  to  be  the  speci- 


124         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


men  discussed  and  described  by  measurement  in 
the  original  publication  and  the  only  adult  speci- 
men mentioned,  is  here  designated  as  lectotype.  P. 
gossijpinus  was  considered  a  valid  species  by  Mus- 
ser  and  Carieton  (1993:  730)  and  Nowak  (1999: 
1361). 

Hesperomys  leucopus  arcticus  Mearns, 

1890 

Bull.  Amer.  Mus.  Nat.  Hist.,  2:  285,  21 

Februaiy. 

Name  preoccupied  by  Hesperomys 

arcticus  Coues,  1 877 

Peromyscus  maniculatus  borealis  Mearns, 

1911 

Proc.  Biol.  Soc.  Washington,  24:  102,  15 

May.  (Replacement  name  for 

Hesperomtjs  leucopus  arcticus  Mearns, 

1890) 

Holotype.  MCZ  5555.  Skin  and  skull.  Adult  male. 
Locality.    Canada:    Northwest  Territories,   Mac- 
Kenzie,  Fort  Simpson,  7  September  1859. 
Collector.  R.  Kennicott.  Original  number  157.  For- 
merly USNM  4531. 

Conclition.  Skin  complete.  Skull  partial  (left  jugal 
and  process  of  squamosal  missing,  right  process  of 
squamosal  missing,  basioccipital  and  palatine  miss- 
ing). Mandible  disarticulated  and  both  angular  pro- 
cess missing. 

Type  Series.  Holot)'pe  only. 

Comments.  P.  m.  borealis  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  672). 

Hesperomys  sonoriensis  nebrascensis 

Coues,  1877 

In  Coues  and  J.  A.  Allen,  Monographs  N. 

Amer.  Rodentia,  U.S.  Geol.  Geogr. 

Survey  Terr.  Rep.  Washington,  11:  79, 

August. 

=  Peromyscus  maniculatus  nebrascensis 

(Coues,  1877).  See  Osgood  (1909:  75). 

Sijntype.  MCZ  5528.  Skin  and  sk-ull. 

Locality.    Nebraska    (now    Wyoming):    (Converse 
County),  Deer  Creek.  19  Januaiy  1860. 
Collector  F.  V.  Hayden.  Original  no.  80.  Formerly 
USNM  4310. 

Condition.  Sldn  complete.  Skull  partial  (posterior 
missing  from  parietal  to  palatine;  both  jugal  and 
process  of  squamosal  broken).  Left  angular  process 
of  mandible  chipped,  right  side  broken  at  cheek 
teeth. 

Type  Series.  1  other  syntype,  present  location  un- 
known, formerly  in  die  USNM. 
Comments.    Hesperomys  sonoriensis   nebrascensis 


Baird,  1858  is  a  nomen  nudum.  Osgood  (1909:  78) 
and  Mearns  (1911:  102)  designated  Deer  Creek, 
Nebraska,  as  the  type  locaHty.  Jones  (1958:  107- 
111)  reviewed  the  systematic  histoiy  oi  nebrascen- 
sis, and  Jones  and  Mursaloglu  (1961:  101-103)  re- 
ported the  discovery  of  one  of  the  two  original  syn- 
types  in  the  MCZ.  The  location  of  the  other  syn- 
tN-pe,  if  it  still  exists,  is  unknown. 

Peromyscus  anastasae  Bangs,  1898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  195,  15 

March. 

=  Peromyscus  gossypinus  anastasae 

Bangs,  1898.  See  Osgood  (1909:  141). 

Holotype.  B7179.  Skin  and  skull.  Adult  female. 
Locality.  (United  States):  Florida,  St.  Johns  Coun- 
ty,   Anastasia    Island,    Point    Romo.    15    February 
1897. 

Collector  O.  Bangs.  Original  number  2. 
Condition.  Sldn  complete,  skull  partial  (premaxilla 
to  frontal  in  fragments).  Right  mandible  missing, 
left  present. 

Type  Series.  There  is  a  small  series  of  paratypes  in 
the  MCZ. 

Comments.  P.  g.  anastasae  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  689). 

Peromyscus  bellus  Bangs,  1896h 

Proc.  Biol.  Soc.  Washington,  10:  137,  28 

December. 

=  Peromyscus  attwateri  ^.  A.  Allen,  1895. 

See  Schmidly  (1973:  125). 

Holott/pe.  B5483.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Indian  Territory 
(  =  Oklahoma),  (Adair  County),  Stilwell.  15  August 
1896. 

Collector  T  Surber.  Original  number  67. 
Condition.  Skin  and  sk-ull  complete.  Mandible  dis- 
articulated. 

Type  Series.  1  paratype;  B5484,  skin  and  skull, 
adult  female. 

Peromyscus  cacabatus  Bangs,  1 902b 
Bull.  Mus.  Comp.  Zool,  39:  29,  April. 
=  Peromyscus  mexicanus  (Saussure, 
1860).  See  Huckaby  (1980:  15). 

Holotype.  MCZ  10225.  Sldn  and  skull.  Adult  female. 
Locality.   Panama:  Chiriqui,  Boquete  (  =  Bajo  Bo- 
quete),  5,000  ft  (1,525  m).  22  April  1901. 
Collector.  W.  W.  Brown,  Jr.  Original  number  331. 
Condition.  Skin  complete.  Skull  partial  (left  jugal 
missing).  Mandible  disarticulated. 
Type  Series.  There  is  a  very  large  series  of  paraty- 
pes in  the  MCZ. 


Type  Specimens  of  Recent  Mammals  •  Helgen  and  McFadden        125 


Peromyscus  canadensis  abietorum  Bangs, 

1896d 

Proc.  Biol.  Soc.  Washington,  10:  49,  9 

March. 

=  Peromyscus  maniculatus  abietorum 

Bangs,  1896.  See  Osgood  (1909:  45). 

Holotype.  B2205.  Skin  and  sknll.  Adult  female. 

Locality.    (Canada):   Nova  Scotia,   James   River.   8 

August  1894. 

Collector.  C.  H.  Goldthw^aite. 

Condition.  Sldn  complete.  Skull  partial  (right  jugal 

missing). 

Type  Series.  There  is  a  series  of  paratypes  in  the 

MCZ. 

Comments.  P.  m.  abietorum  was  retained  as  a  valid 

subspecies  by  Hall  (1981:  670). 

Peromyscus  canadensis  argentatus 

Copeland  and  Church,  1906 

Proc.  Biol.  Soc.  Washington,  19:  122,  6 

September. 

=  Peromyscus  maniculatus  argentatus 

Copeland  and  Church,  1906.  See  Osgood 

(1909:  46). 

Holotype.  MCZ  54627.  Skin  and  skull.  Adult  male. 
Locality.  (Canada):  New  Brunswick,  Grand  Manan 
Island,  Grand  Harbor.  19  September  1905. 
Collector  M.  L.  Church  and  M.  Copeland.  Original 
number  168. 

Condition.  Skin  and  skull  complete. 
Type  Series.  There  is  a  large  series  of  paratypes  in 
the  MCZ. 

Comments.  P.  m.  argentatus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  671). 

Peromyscus  crinitus  scitulus  Bangs,  1899j 

Proc.  New  England  Zool.  Club,  1:  67,  31 

July. 

=  Peromyscus  crinitus  crinitus  (Merriam, 

1891).  See  Osgood  (1909:  229). 

Holotype.  B9175.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Nevada,  Douglas  County, 

Gardnerville.  13  July  1898. 

Collector  W.  W.  Price  and  R  O.  Simons.  Original 

number  1571. 

Condition.  Skin  complete.  Skull  partial  (right  and 

left  jugal  missing). 

Type  Series.   7  paratypes;  B9177-B9180,  B9183- 

B9185;  all  represented  by  skin  and  skull,  4  females 

and  3  males. 


Peromyscus  gossypinus  nigriculus  Bangs, 

1896e 

Proc.  Biol.  Soc.  Washington,  10:  124,  5 

November. 

=  Peromyscus  gossypinus  gossypinus 

(LeConte,  1853).  See  Osgood  (1909:  136). 

Holotype.  B2731.  Sldn  and  skull.  Adult  female. 

Locality.   (United  States):  Louisiana,  Plaquemines 
Parish,  Burbridge.  30  Januaiy  1895. 
Collector  F.  L.  Small.  Original  number  1547/156. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  There  is  a  veiy  large  series  of  paraty- 
pes in  the  MCZ. 

Peromyscus  gossypinus  palmarius  Bangs, 

1896e 

Proc.  Biol.  Soc.  Washington,  10:  124,  5 

November. 

Holotype.  B3224.  Skin  and  skull.  Adult  female. 
Locality.  (United  States):  Florida,  Brevard  County, 
Oak  Lodge  on  East  Peninsula  opposite  Micco.  23 
Februaiy  1895. 
Collector  O.  Bangs. 
Condition.  Skin  and  skull  conaplete. 
Type  Series.  There  is  a  very  large  series  of  paraty- 
pes in  the  MCZ. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  690). 

Peromyscus  insulanus  Bangs,  1898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  196,  15 

March. 

=  Peromyscus  gossypinus  anastasae 

Bangs,  1898.  See  Osgood  (1909:  141). 

Holotype.  B6438.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Georgia,  Camden  Coun- 
ty, Cumberland  Island,  north  end.  10  April  1897. 
Collector  W.  W.  Brown,  Jr.  Original  number  804. 
Condition.  Skin  complete.  Skull  partial  (right  jugal 
missing) . 

Type  Series.  There  is  a  large  series  of  paratypes  in 
the  MCZ. 

Peromyscus  leucopus  ammodytes  Bangs, 

1905a 

Proc.  New  England  Zool.  Club,  4:  14,  28 

Februaiy. 

Holotype.  B828.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Massachusetts,  Barnsta- 
ble County,  Monomoy  Island.  28  December  1893. 
Collector  G.  S.  Miller,  Jr.  and  O.  Bangs.  Original 
number  8. 
Condition.  Skin  and  skull  complete. 


126         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Type  Series.  There  is  a  series  of  paratypes  in  the 

MCZ. 

Comments.  Retained  as  a  vaUd  subspecies  by  Hall 

(1981:  685). 


Peromyscus  leucopus  fusus  Bangs,  1 905a 
Proc.  New  England  Zool.  Club,  4:  13,  28 


Februaiy. 


Holotijpe.  B9737.  Skin  and  skull.  Adult  male. 

Localitij.    (United   States):    Massachusetts,    Dukes 

County,   Martha's  Vineyard,  W.   Tisbury.    17  June 

1899. 

Collector.  O.  Bangs.  Original  number  1. 

Condition.  Skin  and  skull  complete. 

Type  Series.  There  is  a  small  series  of  paratypes  in 

the  MCZ. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 

(19S1:  686). 


Peromyscus  oreas  Bangs,  1 898f 

Proc.  Biol.  Soc.  Washington,  12:  84,  24 

March. 


Holotype.  B3696.  Skin  and  skull.  Adult  female. 

Locality.  Canada:  British  Columbia,  near  boundaiy 

of  Whatcom   County,   Washington,   49th  parallel. 

Mount  Baker  Range,  6,500  ft  (1,983  m).  29  August 

1896. 

Collector  A.  C.  Brooks.  Original  number  745. 

Condition.  Sldn  and  skull  complete. 

Type  Series.  There  is  a  large  series  of  paratypes  in 

the  MCZ. 

Comments.  Considered  a  valid  species  by  Musser 

and  Carleton  (1993:  734)  and  Nowak  (1999;  1361). 

Peromyscus  phasma  Bangs,  1898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  199,  15 

March. 

=  Peromyscus  poloniotus  phasma  Bangs, 

1898.  See  Osgood  (1909:  107). 

Holotype.  B7175.  Skin  and  skull.  Adult  female. 

Locality.  (United  States):  Florida,  St.  Johns  Coun- 
ty, Anastasia  Island,  Point  Romo.  14  Februaiy 
1897. 

Collector  O.  Bangs.  Original  number  3. 
Condition.   Skin  complete.  Skull  partial  (parietal, 
tympantic  bulla,   left  jugal,  and  process  of  squa- 
mosal broken).  Mandible  disarticulated. 
Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  P.  p.  phasma  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  668). 


Peromyscus  subgriseus  arenarius  Bangs, 

1898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  202, 

March. 

Name  preoccupied  by  Peromyscus 

eremicus  arenarius  Mearns,  1 896. 

Peromyscus  subgriseus  baliolus  Bangs, 

1898n 

Science,  n.  sen,  8:  215,  19  August. 

(Replacement  name  for  Peromyscus 

subgriseus  arenarius  Bangs,  1898) 

=  Peromyscus  polionotus  polionotus 

(Wagner,  1843).  See  Osgood  (1909:  104). 

Holotype.  B5925.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Georgia,  Scriven  County, 

Hursman's  Lake  (Savannah  River),  near  Bascom. 

15  December  1896. 

Collector  W.  W.  Brown,  Jr  Original  number  60. 

Condition.  Skin  and  skull  complete. 

Type  Series.  There  is  a  large  series  of  paratypes  in 

the  MCZ. 

Peromyscus  subgriseus  rhoadsi  Bangs, 

1898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  201,  15 

March. 

=  Peromyscus  polionotus  riioadsi  Bangs, 

1898.  See  Osgood  (1909:  107). 

Holotype.  B6980.  Skin  and  skull.  Adult  male. 

Locality.    (United    States):    Florida,    Hillsborough 

County,  head  of  Anclote  River.  23  May  1895. 

Collector.  W.  S.  Dickinson. 

Condition.  Sldn  and  skvdl  complete. 

Type  Series.  There  is  a  series  of  paratypes  in  the 

MCZ. 

Comments.  P.  p.   rhoadsi  was  retained  as  a  valid 

subspecies  by  Hall  (1981:  669). 

Peromyscus  texanus  saturatus  Bangs, 

1897a 

Amer.  Nat.  31:  75,  1  Januaiy. 

=  Peromyscus  maniculatus  saturatus 

Bangs,  1897.  See  Osgood  (1909:  61). 

Holotype.  B2581.  Skin  and  skull.  Adult  male. 

Locality.    Canada:    British   Columbia,   Saturna  Is- 
land. 31  Januaiy  1894. 

Collector  W.  C.  Colt.  Original  number  128. 
Condition.  Skin  and  skull  complete. 
Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  P.  m.  saturatus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  682). 


Type  Specimens  of  Recent  Mammals  •  Helpen  and  McFadden 


127 


Genus  REITHRODONTOMYS  Giglioli, 
1873 

Reithrodontomys  australis  vulcanius 
Bangs,  1902b 

Bull.  Mus.  Comp.  Zool,  39:  38,  April. 
=  Reithrodontomys  sumichrasti  vulcanius 
Bangs,  1902.  See  Hooper  (1952:  83). 

Holotijpe.  MCZ  10281.  Skin  and  skull.  Adult  male. 
Locality.    Panama:   Chiriqui,   Volcan  de   Chiriqui, 
10,300  ft  (3,142  m).  26  May  1901. 
Collector.  W.  W.  Brown,  Jr.  Original  number  500. 
Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Holotype  only. 

Comments.  R.  s.  vulcanius  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  644). 

Reittirodontomys  creper  Bangs,  1902b 
Bull.  Mus.  Comp.  Zool,  39:  39,  April. 

Holotype.  MCZ  10284.  Skin  and  skull.  Adult  female. 
Locality.    Panama:   Chiriqui,   Volcan   de   Chiriqui, 
11,000  ft  (3,355  m).  2  June  1901. 
Collector.  W.  W.  Brown.  Original  number  504. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. Left  lower  incisor  missing. 
Type  Series.  Holotype  only. 

Comments.  Considered  a  valid  species  by  Musser 
and  Carleton  (1993:  740)  and  Nowak  (1999:  1364). 

Reithrodontomys  lecontii  impiger  Bangs, 

1898m 

Proc.  Biol.  Soc.  Washington,  12:  167,  10 

August. 

=  Reithrodontomys  humulis  humulis 

(Audubon  and  Bachman,  1841).  See  Hall 

and  Kelson  (1959:  584). 

Holotype.  B7784.  Skin  and  skull.  Adult  male. 

Locality.    (United   States):   West  Virginia,   Green- 
briar  County,  White  Sulphur  Springs,  2,000  ft  (610 
m).  27  Febmaiy  1898. 
Collector.  T.  Surber.  Original  number  466. 
Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  2  paratypes;  B6931  (not  B6932,  as 
stated  in  the  original  description),  skin  and  skull, 
adult  male;  B7785,  skin  and  skull,  adult  female. 

Genus  SCOT/A/OM/S  Thomas,  1913 

AlKodon  teguina  apricus  Bangs,  1 902b 
Bull.  Mus.  Comp.  Zool,  39:  40,  April. 
=  Scotinomys  teguina  apricus  (Bangs, 
1902).  See  Thomas  (1913:  409). 

Holotype.  MCZ  10236.  Skin  and  skull.  Adult  female. 
Locality.  Panama:  Chiriqui,  Boquete  (  =  Baio  Bo- 
quete),  4,000  ft  (1,220  m).  24  February  1901. 


Collector.  W  W.  Brown,  Jr.  Original  number  192. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  4  paratypes;  MCZ  10234,  skin  and 
skull,  adult  male;  MCZ  10235,  skin,  adult  female; 
MCZ  10237,  skin  and  skull,  adult  female;  MCZ 
10238,  sldn  and  skull,  adult  female. 
Comments.  S.  t.  apricus  was  retained  as  a  valid  sub- 
species by  Hooper  (1972:  21)  and  Hall  (1981:  734). 

AI<odon  xerampelinus  Bangs,  1 902b 
Bull.  Mus.  Comp.  Zool.,  39:  41,  April. 
=  Scotionomys  xerampelinus  (Bangs, 
1902).  See  Thomas  (1913:  409). 

Holotype.  MCZ  10240.  Skin  and  skull.  Adult  male. 
Locality.    Panama:   Chiriqui,   Volcan  de   Chiriqui, 
10,300  ft  (3,142  m).  26  May  1901. 
Collector  W  W  Brown,  Jr.  Original  number  501. 
Condition.  Sldn  complete.  Skull  partial  (right  jugal 
missing).  Mandible  disarticulated. 
Type   Series.   2  paratypes;   MCZ   10239,  skin  and 
skull,  adult  male;  MCZ  10241,  skin  and  skull,  adult 
male. 

Comments.  S.  xerampelinus  was  considered  a  valid 
species  by  Musser  and  Carleton  (1993:  746)  and 
Nowak  (1999:  1355). 

Genus  SIGMODON  Say  and  Ord,  1825 

Sigmodon  austerulus  Bangs,  1 902b 
Bull.  Mus.  Comp.  Zool,  39:  32,  April. 
=  Sigmodon  hispidus  borucae  J.  A.  Allen, 
1897.  See  Hall  and  Kelson  (1959:  672). 

Holotype.  MCZ  10288.  Sldn  and  skull.  Adult  male. 
Locality.    Panama:   Chiiiqui,  Volcan   de   Chiriqui, 
10,000  ft  (3,050  m).  1  June  1901. 
Collector  W  W  Brown,  Jr.  Original  number  503. 
Condition.  Skin  complete.  Skull  partial — according 
to  original  description,  "unfortunately  it  [the  skull] 
was  broken  by  the  trap  directly  across  the  orbits." 
Tijpe  Series.  Holotype  only. 

Comments.  For  comments  regarding  the  type  lo- 
cality, see  Enders  (1953a:  508-509),  who  suggested 
that  a  native  helper  of  W.  W  Brown,  Jr,  transplant- 
ed a  specimen  of  S.  [h.]  borucae,  trapped  during  a 
trip  to  his  home  in  the  lowlands  near  Boruca,  to 
the  top  of  the  Volcan,  "after  Brown  had  promised 
a  bottle  of  wine  to  any  man  who  captured  a  Sig- 
modon  at  that  altitude." 

Sigmodon  hispidus  exsputus  G.  M.  Allen, 

1920b 

J.  Mammal.,  1:  236,  4  December. 

Holotype.  MCZ  18100.  Skin  and  skull.  Adult  male. 
Locality.  (United  States):  Florida,  Monroe  County, 
Big  Pine  Key  16  April  1920. 
Collector  W  S.  Brooks.  Original  number  1936. 


128         Bidletm  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type   Series.    1   paraty][De;    MCZ    18101,   skin   and 
skull,  adult  male. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  737). 


Sigmodon  hispidus  furvus  Bangs,  1 903b 
Bull.  Mus.  Comp.  Zool.,  39:  158,  July. 


Holottjpe.  MCZ  10665.  Skin  and  skull.  Male. 

Locality.  Honduras:  Atlantida,  La  Ceiba.  16  Janu- 
ary 1902. 

Collector.  W.  W.  Brown,  Ji".  Original  number  4. 
Condition.  Skin  and  skull  complete. 
Type  Series.  Holotype  only. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  738).  Sigmodon  hispidus  fei'vidus  Lydekker, 
1904  (p.  34)  was  a  misspelling  and  thus  accidental 
renaming  of  Sigmodon  hispidus  furvtis  Bangs, 
1903. 


Sigmodon  hispidus  spadicipygus  Bangs, 

1898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  192,  15 

March. 


Holotype.  B4477.  Skin  and  skull.  Adult  female. 

Locality.  (United  States):  Florida,  Monroe  County, 
Cape  Sable.  18  April  1895. 

Collector  C.  L.  Brownell.  Original  number  153. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  739). 


Sigmodon  sanctae-martae  Bangs,  1898q 

Proc.  Biol.  Soc.  Washington,  12:  189,  30 

December. 

=  Sigmodon  hispidus  hirsutus  (Burmeister, 

1854).  See  Cabrera  (1961:  508). 


Holotype.  B8105.  Skin  and  skull.  Adult  male. 

Locality.  Colombia:  (Magdalena),  (Santa  Marta 
Mountains),  Pueblo  Viejo  (not  far  from  the  source 
of  Rio  Ancho),  8,000  ft  (2,440  m).  23  March  1898. 
Collector.  W.  W.  Brown,  Jr.  Original  number  73. 
Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type   Series.    1   paratype;   B8250,   skin   and  skull, 
adult  male. 


Genus  THOIVI ASOIVIYS  Coues,  1884 

Oryzomys  (Erioryzomys)  monochromos 

Bangs,  1900b 

Proc.  New  England  Zool.  Club,  1:  97,  23 

February. 

=  Thomasomys  monochromos  (Bangs, 

1900).  See  Gardner  and  Patton  (1976: 

26). 

Holotype.  B8348.  Sldn  and  skull.  Adult  male. 

Locality.  Colombia:  Magdalena,  Santa  Marta 
Mountains,  Paramo  de  Macotania,  11,000  ft  (3,355 
m).  7  March  1899. 

Collector  W.  W.  Brown,  Jr.  Original  number  93. 
Condition.  Skin  complete.  Skull  partial  (right  jugal 
missing).  Majidible  disarticulated. 
Type   Series.    4  paratypes;   B8248,  juvenile   male, 
B8345,  adult  female,  B8346,  adult  female,  B8347, 
adult  male;  all  represented  by  sldn  and  skull. 
Comments.  Considered  a  valid  species  by  Musser 
and  Carleton  (1993:  750)  and  Nowak  (1999:  1362). 
Paramo  de  Macotama  is  in  La  Guajira  (Paynter 
1997:  261). 

Genus  ZYGODONTOMYS  ^.  A.  Allen, 
1897 

Zygodontomys  seorsus  Bangs,  1901b 
Amer.  Nat.,  35:  642,  22  August. 
= Zygodontomys  brevicauda  cherriei  (J.  A. 
Allen,  1895).  See  Voss  (1991:  59). 

Holotype.  B8490.  Skin  and  skull.  Adult  male. 

Locality.  Panama:  Gulf  of  Panama,  San  Miguel  Is- 
land, Isla  del  Rey  5  May  1900. 
Collector  W.  W.  Brown,  Jr  Original  number  147. 
Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Bangs  based  the  description  on  a  se- 
ries of  68  specimens,  most  of  which  are  still  in  the 
MCZ. 

Family  PEDETIDAE  Gray,  1825 

Genus  PEDETES  llliger,  1811 

Pedetes  cafer  taborae  G.  M.  Allen  and 

Loveridge,  1927 

Proc.  Boston  Soc.  Nat.  Hist.,  38:  438,  23 

December. 

=  Pedetes  capensis  (Forster,  1778).  See 

Misonne  (1974:  8). 

Holotype.  MCZ  23080.  Skin  and  skull.  Adult  female. 
Locality.    Tanganyika   Territory    (=Tanzania):    Ta- 
bora,  Unyamwezi,  4,000  ft  (1,220  m).  18  November 
1921. 
Collector  A.  Loveridge.  Original  number  R6915. 


Type  Specimens  of  Recent  Mammals  •  Helgen  and  McFadden        129 


Condition.  Sldn  complete,  with  small  bald  spots  on 
dorsum.  Skull  complete. 
Type  Series.  Holotype  only. 

Family  MYOXIDAE  Gray,  1821 

Genus  GRAPHIURUS  Smuts,  1832 

Aethoglis  hueti  argenteus  G.  M.  Allen, 

1936 

J.  Mammal,  17:  293,  14  August. 

=  Graphiurus  hueti  argenteus  (G.  M.  Allen, 

1936).  See  Hayman  and  Holt  (1940: 

608). 

Holotype.  MCZ  17920.  Skin  and  skull.  Adult  female. 
Locality.  Cameroons  (  =  Cameroon):  (Kribi),  Lolo- 
dorf.  17  March  1911. 

Collector.  G.  Schwab.  Original  number  1. 
Condition.  Sldn  and  skull  complete. 
Type   Series.   2  paratypes;   MCZ  17607,  sldn  and 
skull,  adult  female;  and  a  specimen  in  the  USNM, 
number  125434. 

Comments.   Rosevear  considered  G.   h.   argenteus 
"possibly  valid  as  a  race"  (1969:  501). 

Claviglis  soleatus  collaris  G.  M.  Allen  and 

Loveridge,  1933 

Bull.  Mus.  Comp.  Zool.,  75:  122, 

Februan'. 

=  Graphiurus  lorraineus  Dollman,  1910. 

See  Holden  (1993:  764). 

Holotype.  MCZ  26373.  Skin  and  skull.  Adult  female. 
Locality.  Tanganyika  Territory  (=Tanzania):  (Irin- 
ga),  north  end  Lake  Nyasa,  Uldnga  Mountains,  Ma- 
dehani,  7,000  ft  (2,135  m).  24  Februaiy  1930. 
Collector.  A.  Loveridge. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type   Series.   2  paratypes;   MCZ  26374,  skin  and 
skull,  male;  MCZ  26375,  skin  and  skull,  female. 

Graphiurus  microtis  griseus  G.  M.  Allen, 

1912a 

Bull.  Mus.  Comp.  Zool,  54:  440,  April. 

=  Grapfiiurus  murinus  griseus  G.  M.  Allen, 

1912.  See  Hayman  and  Holt  (1940:  610). 

Holotype.  MCZ  8244.  Skin  and  skull.  Adult  male. 
Locality.   British  East  Africa  (  =  Kenya):  northern 
Guaso  Nyiro  River  (  =  Ewaso  Ngiro),  4,000  ft  (1,220 
m).  25  July  1909. 

Collector  G.  M.  Allen.  Original  number  51. 
Condition.  Skin  and  skull  complete. 
Type  Series.  2  paratypes;  MCZ  8248,  skin  and  skull, 
male;  MCZ  8249,  skin  and  skaiU,  adult  female. 


Grapiiiurus  scfiwabi  G.  M.  Allen,  1912a 
Bull.  Mus.  Comp.  Zool.,  54:  441,  April. 
=  Grapiiiurus  surdus  Dollman,  1912.  See 
Holden  (1993:  765). 


Holotype.  MCZ  8607.  Skin  and  skull.  Juvenile. 
Locality.  Cameroun  (  =  Cameroon),  Kribi.  1911. 
Collector  G.  Schwab. 
Condition.  Skin  and  skull  complete. 
Type  Series.  Holotype  only. 

Family  BATHYERGIDAE  Waterhouse, 
1841 


Genus  CRYPTOIVIYS  Gmy,  1864 

Cryptomys  tiottentotus  occlusus  G.  M. 

Allen  and  Loveridge,  1933 

Bull.  Mus.  Comp.  Zoo).,  75:  125, 

Februaiy. 

=  Cryptomys  tiottentotus  wfiytei  (Thomas, 

1897).  See  Honeycutt  et  al.  (1991:  51). 

Holotype.  MCZ  26557.  Skin  and  skull.  Adult  male. 
Locality.  Tanganyika  Territoiy  (=Tanzania):  (Irin- 
ga),  Ugungwe  Mountains,  Kigogo,  6,000  ft  (1,830 
m).  18  January  1930. 
Collector.  A.  Loveridge. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.   15  paratypes;  MCZ  26558-26572,  all 
represented  by  skin  and  skull. 

Genus  HETEROCEPHALUS  Ruppell, 
1842 

l-ieterocepiiaius  stygius  G.  M.  Allen, 

1912a 

Bull.  Mus.  Comp.  Zool,  54:  444,  April. 

=  l-leterocepiiaius  giaber  Ruppell,  1842. 

See  HoUister  (1919:  160). 


Holotype.  MCZ  12470.  Body  in  alcohol,  skull  e.xtract- 
ed.  Adult  female. 

Locality.    British  East  Africa  (  =  Kenya):  northern 
Guaso  Nyiro  (  =  Ewaso  Ngiro),  Neumann's  Boma.  6 
August  1909. 
Collector  G.  M.  Allen. 

Condition.    Alcoholic.    Skull   complete.    Mandible 
disarticulated. 
Type  Series.  Holotype  only. 


130         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Family  ERITHIZONTIDAE  Bonaparte, 
1845 

Genus  ERITHIZONF.  Cuvier,  1822 

Erethizon  dorsatus  picinus  Bangs,  1900c 

Proc.  New  England  Zool.  Club,  2:  37,  20 

September. 

=  Erithizon  dorsatum  dorsatum  (Linnaeus, 

1758).  See  Hai-per  (1961:  90). 

Holotype.  B8839.  Sldn  and  skull.  Adult  male. 

Locality.  (Canada):  Labrador,  Strait  of  Belle  Isle, 
Lance  (  =  Lanse)  au  Loup.  16  Februaiy  1899. 
Collector.  E.  Doane. 
Condition.  Skin  and  skull  complete. 
Type  Series.   1.5  paratypes;  B8832-B8838,  B8840- 
B8847;  all  represented  by  skin  and  skull,  8  females 
and  7  males.   Bangs  had  also  a  skull  of  picimis 
(B8831)  from  Black  Bay,  Labrador,  at  the  time  of 
description,  not  mentioned  in  the  original  publi- 
cation. 

Family  DASYPROCTIDAE  Gray,  1825 

Genus  DASYPROCTA  llliger,  1811 

Dasyprocta  callida  Bangs,  1 901  b 
Amer.  Nat.,  35:  635,  22  August. 
=  Dasyprocta  punctata  callida  Bangs, 
1901.  See  Kellogg  (1946:  59). 

Holotype.  B8443.  Skin  and  skull.  Adult  male. 

Locality.  Panama:  (Panama),  San  Miguel  Island.  8 

May  1900. 

Collector.  W.  W.  Brown,  Jr  Original  number  171. 

Condition.  Skin  and  skull  complete. 

Type  Series.  5  paratypes;  B8442,  B8444-B8447;  all 

represented  by  skin  and  skull,   3  females  and  2 

males. 

Comments.   D.  p.  callida  was  retained  as  a  valid 

subspecies  by  Hall  (1981:  860). 

Dasyprocta  colombiana  Bangs,  1 898k 

Proc.  Biol.  Soc.  Washington,  12:  163,  10 

August. 

=  Dasyprocta  punctata  colombiana  (Gray, 

1898).  See  Cabrera  (1961:  589). 

Holotype.  B8008.  Skin  and  skull.  Adult  female. 

Locality.    Colombia:    Magdalena,    Santa   Marta.   6 
January  1898. 

Collector.  W.  W.  Brown,  Jr.  Original  number  37. 
Condition.  Sldn  and  skull  complete. 
Type  Series.  1  paratype;  B8113,  skin  and  skull,  ju- 
venile male. 

Comments.  D.  p.  colombiana  was  retained  as  a  val- 
id subspecies  by  Cabrera  (1961:  .589). 


Dasyprocta  noblei  G.  M.  Allen,  1914e 

Proc.  New  England  Zool.  Club,  5:  69,  7 

October. 

=  Dasyprocta  leporina  noblei  G.  M.  Allen, 

1914.  See  Woods  (1993:  781). 

Holotype.  MCZ  15936.  Sldn  and  skull  (and  atias). 
Adult  female. 

Locality.   Guadeloupe  Island:  Goyave.  22  August 
1914. 

Collector  G.  K.  Noble. 

Condition.  Sldn  complete.  Skull  partial  (bone  miss- 
ing from  nasal  and  frontal). 

Type   Series.    1   paratype;    MCZ   15937,   sldn   and 
skull,  subadult  female. 

Dasyprocta  punctata  nuchalis  Goldman, 

1917 

Proc.  Biol.  Soc.  Washington,  30:  113,  23 

May. 

Holotype.  MCZ  1008  L  Skin  and  skull.  Adult  female. 
Locality.  Panama:  Chiriqui,  Divala.  30  November 
1900. 

Collector  W.  W.  Brown,  Jr.  Original  number  17. 
Condition.  Skin  and  skull  complete. 
Type   Series.   4  paratypes;   MCZ  10080,  skin  and 
skull,   adult  male;   10084,   skull;   10175,   skin  and 
skull,   adult  female;    10176,   sldn  and  skull,   adult 
male. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  860). 

Family  AGOUTIDAE  Gray,  1821 

Genus  AGOUTI  Lacepede,  1799 

Agouti  paca  virgatus  Bangs,  1902b 
Bull.  Mus.  Comp.  Zool.,  39:  47,  April. 

Holotype.  MCZ  10079.  Sldn  and  skull.  Adult  male. 
Locality.  Panama:  Chiriqui,  Divala.  16  December 
1900. 

Collector  W.  W.  Brown,  Jr  Original  number  21. 
Condition.  Sldn  and  skull  complete. 
Type  Series.  Holotype  only. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  8.58)  and  Perez  (1992:  1). 

Family  OCTODONTIDAE  Waterhouse, 

1839 

Genus  TYMPANOCTOMYS  Yepes,  1941 

Octomys  barrerae  Lawrence,  1941 
Proc.  New  England  Zool.  Club,  18:  43, 
28  Januaiy. 

=  tympanoctomys  barrerae  (Lawrence, 
1941).  See  Yepes  (1942:  75). 

Holotype.  MCZ  39716.  Skin  and  skull.  Aduk  male. 
Locality.   Argentina:    Mendoza  Province,   La  Paz. 
April  1939. 


Type  Specimens  of  Recent  Mammals  •  Helpen  and  McFadden 


131 


Collector.   J.   M.   de  la  Barrera.   Original  number 

A31. 

Condition.  Sldn  and  skull  complete. 

Type  Series.  Holotype  only. 

Comments.  Type  species  of  Tyinpanoctomys  Yepes, 

1941.  T.  barrerae  was  considerd  a  valid  species  by 

Woods  (1993:  789)  and  Nowak  (1999:  1683). 

Family  ECHIMYIDAE  Gray,  1825 

Genus  BOROMYS  M\\\er,  1916 

Boromys  torrei  G.  M.  Allen,  1917a 
Bull.  Mus.  Comp.  Zool.,  61:  6,  Januaiy. 

Holotype.  VP  9601.  Palate. 

Locality.  Cuba:  Matanzas  Province,  cave  in  Sierra 
de  Hato  Nuevo. 
Collector  C.  de  la  Torre. 

Condition.  Condition  as  described  in  original — 
"palate  with  root  of  right  z}'gomatic  arch,  pni^  and 
alveolar  row  of  right  side,  i)i'  and  posterior  part  of 
alveolar  row  of  left  side." 

Type  Series.  Paratype  material  consists  of  8  lower 
jaws  and  2  separate  lower  molars;  in  the  MCZ. 
Comments.  Considered  a  valid  species  by  Woods 
(1993:  799)  and  Nowak  (1999:  1703)  but  almost 
certainly  extinct.  This  specimen  is  stored  in  the 
Vertebrate  Paleontology  Department  of  tlie  MCZ. 

Genus  DIPLOMYS  Thomas,  1916 

Loncheres  labilis  Bangs,  1901b 
Amer.  Nat.,  35:  638,  22  August. 
=  Diplomys  labilis  (Bangs,  1901).  See 
Thomas  (1916:  296). 

Holotype.  B8480.  Skin  and  skull.  Adult  male. 

Locality.  Panama:  (Panama),  Gulf  of  Panama,  San 
Miguel  Island.  26  April  1900. 

Collector.  W  W  Brown,  Jr.  Original  ninuber  103. 
Condition.  Skin  and  skull  complete. 
Type  Series.  There  is  a  small  series  of  paratypes  in 
the  MCZ  and  FMNH. 

Comments.  D.  labilis  was  considered  a  valid  spe- 
cies by  Woods  (1993:  791)  and  Nowak  (1999: 
1695). 

Genus  PROECHIMYS  ^.  A.  Allen,  1899 

Proechimys  burrus  Bangs,  1901b 
Amer.  Nat.,  35:  640,  22  August. 
=  Proechimys  semispinosus  burrus  Bangs, 
1901.  See  Goldman  (1920:  120). 

Holotype.  B8458.  Skin  and  skull.  Adult  male. 

Locality.  Panama:  (Panama),  Gulf  of  Panama,  San 
Miguel  Island.  30  April  1900. 
Collector  WW.  Bro\\ai,  Jr.  Original  number  130. 
Condition.  Skin  and  skull  complete.  Bight  dentary 
broken  at  pnij. 


Type  Series.  There  is  a  large  series  of  paratypes  in 
the  MCZ. 

Comments.  P.  s.  burnis  was  retained  as  a  valid  sub- 
species bv  Hall  (1981:  873). 

Proechimys  gorgonae  Bangs,  1905b 
Bull.  Mus.  Comp.  Zool.,  46:  89,  June. 

Holotype.  MCZ  10828.  Sldn  and  skull.  Adult  male. 
Locality.    Colombia:    (Cauca),    Gorgona   Island.   2 
July  1904. 

Collector  W.  W  Brown,  Jr.  Original  number  25. 
Condition.  Skin  and  skull  complete. 
Tijpe  Series.   6  paratypes;  MCZ  10829-10834,  all 
represented   by   skin   and   skull,    1    female   and  5 
males.    MCZ    10830   and    10834   are   now  in   the 
FMNH. 

Comments.  Treated  as  a  separate  species  by  Woods 
(1993:  796)  but  included  in  P.  cayennensis  by  No- 
wak (1999:  1689). 

Proechimys  guyannensis  hyleae  Moojen, 

1948 

Univ.  Kansas  Publ.  Mus.  Nat.  Hist.,  1: 

361,  10  December. 

=  Proechimys  cayennensis  hyleae  Moojen, 

1948.  See  comments. 

Holotype.  MCZ  30887.  Skin  and  skull.  Adult  male. 
Locality.  Brazil:  Para,  Porto  de  Moz,  Rio  Tapajoz 
(=Tapajos),  Tauari  (=Tauaiy),  87  km  south  of  San- 
tarem.  19  January  1934. 

Collector  A.  M.  Olalla.  Original  number  7288. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  20  paratypes  (19  at  MCZ  and  1  at 
FMNH),  represented  by  sldn  and  skull. 
Comments.  Retained  as  a  valid  subspecies  by  Ca- 
brera (1961:  521).  The  use  of  the  specific  name 
cayennensis  over  guyannensis  follows  Woods 
(1993:  795). 

Family  CAPROMYIDAE  Smith,  1842 

Genus  CAPROMYS  Desmarest,  1822 

Capromys  pilorides  relictus  G.  M.  Allen, 

1911a 

Bull.  Mus.  Comp.  Zool,  54:  207,  July. 

Holotype.  MCZ  10996.  Skin  and  skull.  Adult  male. 
Locality.  Cuba:  Isle  of  Pines  (  =  Isla  de  Pinos),  Nue- 
va  Gerona,  Casas  Mountains.  10  March  1902 
Collector.  W.  R.  Zappey. 

Condition.  Sldn  complete.  Skull  partial  (bone  miss- 
ing from  palatine  to  occipital). 

Type  Series.  1  paratype;  MCZ  10997,  sldn  and 
skull. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  863). 


132         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Genus  GEOCAPROMYS  Chapman,  1901 

Geocapromys  cubanus  G.  M.  Allen,  1917a 
Bull.  Mus.  Comp.  Zool.  61:  9,  Januaiy. 
=  Geocapromys  columbianus  (Chapman, 
1892).  See  G.  M.  Allen  (1918b:  145). 

Holotype.  VP  9602.  Part  of  right  lower  ramus.  Juve- 
nile. 

Locality.  Cuba:  Matanzas  Province,  cave  in  Sierra 
de  Hato  Neuvo. 
Collector.  C.  de  la  Torre. 

Condition.   Portion  of  ramus  includes  incisor  and 
three  anterior  cheekteeth. 

Type  Series.  Allen  mentions  in  the  original  descrip- 
tion, "Five  palates  with  teeth,  about  15  jaw  frag- 
ments mostly  with  teeth,  and  numerous  other  frag- 
ments." This  material  is  in  the  MCZ. 
Comments.  The  type  material  o{  cubanus  is  stored 
in  the  Vertebrate  Paleontology  Department  of  the 
MCZ. 

Geocapromys  ingrahami  abaconis 
Lawrence,  1934 

Occas.  Pap.  Boston  See.  Nat.  Hist.,  8: 
190,  7  November. 

Holotype.  VP  2108.  Left  lower  ramus.  Adult. 

Locality.   Bahamas:  Great  Abaco  Island,   Hole  in 
the  Wall,  Imperial  Lighthouse  Caves.  1934. 
Collector  F.  Rainey. 
Condition.  Ramus  intact. 

Type  Series.  The  parats'pe  series  is  a  collection  of 
cranial  and  mandibular  fragments,  described  in  the 
original  description  and  housed  in  the  MCZ. 
Comments.  G.  i.  abaconis  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  866)  under  the  genus 
Capromys.  Woods  (1993:  800)  employed  the  genus 
Geocapromys.  G.  i.  abaconis  is  extinct.  The  type 
material  of  abaconis  is  stored  in  the  Vertebrate  Pa- 
leontology Department  of  the  MCZ. 

Geocapromys  ingrahami  irrectus 
Lawrence,  1934 

Occas.  Pap.  Boston  Soc.  Nat.  Hist.,  8: 
190,  7  November. 

Holotype.  VP  2107.  Right  lower  ramus.  Adult. 

Locality.   Bahamas:  Crooked  Island,  Gordon  Hill 
Caves,  "Burial  Cave  No.  1."  1934. 
Collector.  F.  Rainey. 
Condition.  Ramus  intact. 

Tijpe  Series.  The  paratype  series  is  a  collection  of 
cranial  and  mandibular  fragments,  described  in  the 
original  description  and  housed  in  the  MCZ. 
Comments.  G.  i.  irrectus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  866)  vmder  the  genus 
Capromys.  Woods  (1993;  800)  employed  tlie  genus 
Geocapromys.  G.  i.  irrectus  is  extinct.  The  type  ma- 


terial of  irrectus  is  stored  in  the  Vertebrate  Pale- 
ontology Department  of  tlie  MCZ. 

Genus  MESOCAPROMYS  Marona,  1970 

Capromys  nana  G.  M.  Allen,  1917b 

Proc.  New  England  Zool.  Club,  6:  54,  28 

March. 

=  Mesocapromys  nanus  (G.  M.  Allen, 

1917).  See  Kratochvil  et  al.  (1978:  15). 


Holotype.  VP  9864.  Right  mandible. 

Locality.  Cuba:  Matanzas,  cave  in  Sierra  de  Hato 
Nuevo.  March  1917. 
Collector.  T.  Barbour. 

Condition.  Mandible  partial — coronoid  and  angu- 
lar processes  broken  off. 

Tijpe  Series.  A  series  of  paratype  material  is  stored 
at  the  MCZ. 

Comments.  G.  M.  Allen  supplemented  his  descrip- 
tion of  Capromys  nanus  after  Thomas  Barbour 
provided  him  with  a  freshly  killed  specimen  in 
1918  (1918b:  140-145).  Type  species  of  the  sub- 
genus Paracapromys  Kratochvil,  Rodriguez,  and 
Baiois,  1978.  M.  nanus  was  considered  a  valid  spe- 
cies by  Woods  (1993:  801)  and  Nowak  (1999: 
1703).  The  type  series  is  stored  in  the  Vertebrate 
Paleontology  Department  of  the  MCZ. 

Genus  PLAGIODONTA  F.  Cuvier,  1836 


Plagiodonta  araeum  Ray,  1 964 
Breviora,  Mus.  Comp.  Zool.,  203:  2,  10 
April. 

Holotype.  VP  7675.  Left  upper  cheektooth,  originally 
described  as  "almost  certainly  the  fourth  (decidu- 
ous) premolar." 

Locality.  Dominican  Republic:  San  Rafael  Prov- 
ince, Hondo  Valle  Municipality,  unnamed  cave  2 
km  southeast  of  Rancho  de  La  Guardia.  April  1963. 
Collector  R.  Allen  and  C.  E.  Ray. 
Condition.  Condition  of  tooth  as  originally  de- 
scribed— "damaged  slighdy  along  the  posterolabial 
wall." 

Type  Series.  Holotype  only. 

Comments.  Considered  a  valid  species  by  Woods 
(1993:  804)  and  Nowak  (1999:  1708).  Complete 
cranial  and  dentaiy  material  of  araeum  has  been 
collected  in  Haiti  and  deposited  in  the  Florida  Mu- 
seum of  Natural  Histoiy  (Woods  1993:  804).  This 
species  is  extinct.  This  specimen  is  stored  in  the 
Vertebrate  Paleontology  Department  of  the  MCZ. 


Type  Specimens  of  Recent  Mammals  •  Helaen  and  McFadder 


133 


Order  LAGOMORPHA  Brandt,  1855 

Family  OCHOTONIDAE  Thomas,  1897 

Genus  OCHOTONA  Link,  1795 

Ochotona  cuppes  Bangs,  1899g 

Proc.  New  England  Zool.  Club,  1:  40,  5 

June. 

=  Ochotona  princeps  cuppes  Bangs,  1899. 

See  A.  H.  Howell  (1924:  27). 

Holotype.  B7389.  Skin  and  skull.  Adult  male. 

Locality.  (Canada):  British  Columbia,  Gold  Range, 
Monishee  Divide,  4,000  ft  (1,220  m).  2  August 
1897. 

Collector.  A.  C.  Brooks.  Original  number  10.30. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  3  paratypes;  MCZ  7390-7392;  all  rep- 
resented by  sldn  and  skull,  all  female.  MCZ  7390 
is  now  at  FMNH. 

Comments.  O.  p.  cuppes  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  289)  and  Smith  and  Wes- 
ton (1990:  1). 

Ochotona  saxatilis  Bangs,  1 899g 

Proc.  New  England  Zool.  Club,  1:  41,  5 

June. 

=  Ochotona  princeps  saxatilis  Bangs, 

1899.  See  A.  H.  Howell  (1924:  27). 

Holotype.  MCZ  2703.  Skin.  Adult  male. 

Locality.  (United  States):  Colorado,  Park  County, 
Snowy  Range,  near  Mount  Lincoln,  Montgomery. 
27  July  1871. 

Collector.  J.  A.  Allen,  Rocky  Mountain  Expedition. 
Original  number  945. 
Condition.  Skin  complete. 

Type  Series.  Large  paratype  series;  MCZ  209,  243- 
263,  skulls;  B41  and  B42,  each  represented  by  sldn 
and  skull;  MCZ  2673-2703,  skins. 
Comments.  O.  p.  saxatilis  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  291)  and  Smith  and  Wes- 
ton (1990:  1). 

Family  LEPORIDAE  Fischer  de  Waldheim, 
1817 

Genus  LEPL/S  Linnaeus,  1758 

Lepus  (Macrotolagus)  alleni  palitans 

Bangs,  1900a 

Proc.  New  England  Zool.  Club,  1:  85,  23 

February. 

Holotype.  B9096.  Skin  and  skull.  Adult  female. 

Locality.  Mexico:  Sinaloa,  Aguacaliente  (about  40 
miles  [64.4  km]  southeast  of  Mazatlan).  7  August 
1897. 


Collector  P.  O.  Simons.  Original  number  157. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated; left  ramus  chipped;  two  holes  in  right 
ramus. 

Type  Series.  1  paratype;  B9097,  skin,  adult  male. 
Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  332)  and  Best  and  Heniy  (1993:  1). 

Lepus  americanus  struthopus  Bangs, 

1898e 

Proc.  Biol.  Soc.  Washington,  12:  81,  24 

March. 

Holotype.  B2025.  Skin  and  skull.  Adult  female. 

Locality.   (Canada):  Nova  Scotia,  Digby.  4  August 

1894. 

Collector  O.  Bangs.  Original  number  9. 

Condition.  Skin  and  skull  complete. 

Type  Series.  There  is  a  series  of  paratypes  in  the 

MCZ. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 

(1981:  318). 

Lepus  arcticus  bangsii  Rhoads,  1896a 
Amer.  Nat.,  30:  236,  6  March. 

Holotype.  B3752.  Skin  and  skull.  Adult  female. 

Locality.  (Canada):  Newfoundland,  Codroy.  3  Au- 
gust 1895. 

Collector  E.  Doane.  Original  number  1. 
Condition.  Sldn  and  skull  complete. 
Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  320). 

Lepus  bairdii  Hayden,  1869 

Amer.  Nat.,  3:  115,  May. 

=  Lepus  americanus  bairdii  Hayden,  1869. 

See  J.  A.  Allen  (1875:  431). 

Syntype.  MCZ  5501.  Sldn  and  skull.  Adult. 

Locality.  (United  States):  Wyoming,  (Fremont 
County),  summit  of  Wind  River  Mountains,  near 
Fremont  Peak.  4  June  1860. 

Collector.  F.  V.  Hayden.  Original  number  90.  For- 
merly USNM  4264. 
Condition.  Sldn  and  skull  complete. 
Tijpe  Series.  2  other  adult  syn types;  USNM  4262/ 
38001,  skin  and  skull,  male;  USNM  4263/4273, 
sldn  and  skidl;  see  Poole  and  Schantz  (1942:  210). 
Comments.  No  type  is  designated  in  the  original 
description,  which  is  based  on  6  individuals  col- 
lected by  Hayden — 3  adults  and  3  juveniles,  origi- 
nally deposited  in  the  USNM  (Poole  and  Schantz 
1942:  210). 


134         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Lepus  bairdi  cascadensis  Nelson,  1907 

Proc.  Biol.  Soc.  Washington,  20:  87,  11 

December. 

=  Lepus  americanus  cascadensis  Nelson, 

1907.  See  Racey  and  Cowan  (1936: 

HIS). 

Holotype.  B1886.  Skin  and  skull.  Adult  male. 

Locality.    Canada:   British  Columbia,   near  Hope, 

Roab's  Ranch.  12  June  1874. 

Collector.  W.  C.  Colt.  Original  number  476. 

Condition.  Sldn  complete.  Skull  slightly  damaged 

(supraoccipital  and  occipital  chipped). 

Type  Series.  Holotype  only. 

Comments.  L.  a.  cascadensis  was  retained  as  a  valid 

subspecies  by  Hall  (1981:  315). 

Genus  Sy/.WMGL/S  Gray,  1867 

Lepus  (Tapeti)  incitatus  Bangs,  1901b 
Amer.  Nat.,  35:  633,  22  August. 
=  Sylvilagus  brasiliensis  incitatus  (Bangs, 
1901).  See  Hershkovitz  (1950:  352). 

Holotype.  B8441.  Skin  and  skull.  Adult  female. 

Locality.  Panama:  (Panama),  Bay  of  Panama,  San 

Miguel  Island.  30  April  1900. 

Collector  W.  W.  Browi:i,  Jr.  Original  number  127 

Condition.  Skin  and  skull  complete. 

Type  Series.  Holotype  only. 

Comments.  S.  b.  incitatus  was  retained  as  a  valid 

subspecies  by  Hall  (1981:  296). 

Lepus  paludicola  Miller  and  Bangs,  1894 

Proc.  Biol.  Soc.  Washington,  9:  105,  9 

June. 

=  Sylvilagus  palustn's  paludicola  (Miller 

and  Bangs,  1894).  See  Nelson  (1909: 

269). 

Holotype.  B1451.  Sldn  and  skull.  Adult  female. 

Locality.  (United  States):  Florida,  (Citrus  County), 

Fort  Island,  near  Crystal  River.  28  Januaiy  1894. 

Collector  F.  L.  Small.  Original  number  11.50. 

Condition.  Skin  and  skull  complete. 

Type  Series.   3  paratypes;  B1452,  skin  and  skull, 

adult   male;    B1453,    sldn    and   skull,    adult   male; 

B1454,  skin  and  skull,  adult  female. 

Connnents.  S.  p.  paludicola  was  retained  as  a  valid 

subspecies  by  Hall  (1981:  299)  and  Chapman  and 

Willner  (1981:  1). 

Lepus  sylvaticus  alacer  Bangs,  1 896h 

Proc.  Biol.  Soc.  Washington,  10:  136,  28 

December. 

=  Sylvilagus  floridanus  alacer  (Bangs, 

1896).  See  Lyon  (1904:  336). 

Holotype.  B5480.  Sldn  and  skull.  Adult  female. 

Locality.       (United      States):      Indian      Territory 


(=Oklahoma),  (Adair  County),  Stilwell.  14  August 
1896. 

Collector.  T.  Siu-ber.  Original  number  65. 
Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  3  paratypes;  B1677,  skin  and  skull, 
B1678,  skin  and  skull,  adult  male;  B5481,  skin  and 
skull,  adult  female. 

Comments.  S.  f.  alacer  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  301)  and  Chapman  et  al. 
(1980:  1). 

Lepus  sylvaticus  transitionalis  Bangs, 

1895a 

Proc.  Boston  Soc.  Nat.  Hist.,  26:  405,  31 

January. 

=  Sylvilagus  transitionalis  (Bangs,  1895). 

See  Nelson  (1909:  195). 

Holottjpe.  B2407.  Skin  and  skull.  Adult  female. 

Locality.  (United  States):  Connecticut,  New  Lon- 
don County,  Libeity  Hill.  6  November  1894. 
Collector  O.  Bangs. 

Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  S.  transitionalis  was  considered  a  valid 
species  by  Hoffman  (1993:  827)  and  Nowak  (1999: 
1727). 

Order  SCANDENTIA  Wagner,  1855 

Family  TUPAIIDAE  Gray,  1825 

Genus  TUPAIA  Raffles,  1822 

Tana  tana  griswoldi  CooMge,  1938 

Proc.  New  England  Zool.  Club,  17:  45,  6 

May. 

=  Tupaia  tana  paitana  (Lyon,  1913).  See 

Medway  (1965:  76). 

Holotype.    MCZ   36416.    Sldn,    skaill,    and   baculum. 
Subadult  male. 

Locality.  (Malaysia):  British  North  Borneo  (  =  Sa- 
bah).  Mount  Kinabalu,  Kenokok  River,  Kiau 
(  =  Kampong  Kiau),  3,300  ft  (1,007  m).  9  August 
1937. 

Collector  J.  A.  Griswold,  Jr.,  Asiatic  Primate  Ex- 
pedition. Original  number  684. 
Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 
Type  Series.  Holotype  only. 


Type  Specimens  of  Recent  Mammals  •  Helpen  and  McFadden 


135 


Order  PRIMATES  Linnaeus,  1758 

Family  INDRIDAE  Burnett,  1828 

Genus  PROPITHECUS  BenneW,  1832 

Propithecus  perrieri  Lavauden,  1931 

Compt.  Rend.  Acad.  Sci.  Paris,  193:  77,  6 

July. 

=  Propithecus  diadema  perrieri  Lavauden, 

1931.  See  W.  C.  O.  Hill  (1953:  568). 

Holotype.    MCZ  44857.   Skin,  skull,  and  postcranial 
skeleton.  Adult  male. 

Locality.  Malagasy  Republic  (Madagascar):  (Toli- 
aiy),  Ifotaka,  vicinity  of  Ambovombe.  Probably 
1928. 

Collector.  M.  Perrier.  Original  number  1.  Grandi- 
dier  Collection. 

Conclition.  Skin  and  skull  complete.  Postcranial 
skeleton  partial  (right  humerus  broken,  foot  bones 
left  in  sldn). 

Type  Series.  3  paratypes;  MCZ  44858,  skin,  skull, 
and  postcranial  skeleton,  adult  female;  MCZ 
44859,  skin  and  skull,  juvenile;  MCZ  46001,  skin 
and  leg  bones,  adult. 

Comments.  P.  d.  perrieri  was  retained  as  a  valid 
subspecies  by  Tattersall  (1982:  103). 

Family  DAUBENTONIIDAE  Gray,  1863 

Genus  DAUBENTONIA  E.  Geoffrey,  1795 

Ctieiromys  madagascariensis  lanigerG. 

Grandidier,  1930d 

Bull.  Acad.  Malgache,  n.  sen,  11:  106 

(for  1928). 

=  Daubentonia  madagascariensis  (Gmelin, 

1788).  See  G.  M.  Allen  (1939:  134). 

Holotype.   MCZ  45947,  Skull  and  postcranial  skele- 
ton. Adult  male. 

Locality.  Madagascar:  Forest  of  the  East, 
Collector.    Received   by   G,    Grandidier  from   the 
Academic  Malgache,  1927-28. 
Condition.  Skull  complete.  Postcranial  skeleton  in- 
complete (hands  and  feet  missing,  presumably  in- 
tact within  mounted  skin). 
Type  Series.  Holotype  only. 

Comments.  The  mounted  skin  of  the  holotype  is  in 
the  collection  of  the  Academic  Malgache  (Anta- 
nanarivo Museum).  Grandidier  named  laniger  as  a 
new  subspecies  on  account  of  its  woolly  pelage,  but 
it  is  probably  just  a  molting  individual  (Schwarz 
1931:  428). 


Family  GALAGONIDAE  Gray,  1825 

Genus  GALAGOIDES  A.  Smith,  1833 

Galago  demidovii  orinus  Lawrence  and 

Wasliburn,  1936 

Occ.  Pap.  Boston  Soc.  Nat.  Hist.,  8:  259, 

8  January. 

=  Galagoides  orinus  (Lawrence  and 

Washburn,  1936).  See  Honess  (1996:  58). 

Holotype.  MCZ  22453,  Skin  and  skull.  Adult  male. 
Locality.   Tanganyika  Territoiy  (=Tanzania):  Mor- 
ogoro,  Uluguru  Mountains,  Bagilo,  5,000  ft  (1,525 
m).  17  September  1926, 
Collector  A.  Love  ridge. 
Condition.  Skin  and  skull  complete. 
Type  Series.  Holotype  only. 

Family  GEBIDAE  Bonaparte,  1831 

Genus  ALOUATTA  Lacepede,  1799 

Alouatta  palliata  luctuosa  Lawrence, 

1933b 

Bull.  Mus.  Comp.  Zool.,  75:  337, 

November. 

= Alouatta  pigra  Lawrence,  1933.  See 

Smith  (1970:  363). 

Holotype.  MCZ  24059.  Skin  and  skTill.  Adult  male. 
Locality.    British  Honduras  (  =  Belize):   Cayo  Dis- 
trict, Mount  Cow.  12  April  1928. 
Collector.  O.  L.  Austin,  Jr.  Original  number  723, 
Condition.  Skin  and  skull  complete, 
Tijpe  Series.  HolotyjDe  only, 

Alouatta  palliata  trabeata  Lawrence, 

1933b 

Bull.  Mus.  Comp.  Zool.,  75:  328, 

November. 

Holotype.  MCZ  29545.  Skin  and  skull.  Adult  male. 
Locality.    Panama:    Herrera    Province,    Capina, 
March  1933. 

Collector  T  Barboui\  Original  number  4, 
Condition.  Skin  and  skull  complete, 
Tijpe  Series.  Lawrence  mentions  examining  19 
specimens  of  trabeata  and  lists  them  by  locality 
(1933:  330),  9  of  these  are  in  the  MCZ;  other  tlian 
the  holotype,  they  are  MCZ  27784,  27785,  28735, 
28736,  29543,  29544,  29546,  and  29548,  1  paraty- 
pe,  MCZ  29547,  has  been  exchanged  to  the  Museu 
Palista,  Sao  Paolo,  Brazil. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  263), 


136         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Genus  CEBUS  Erxleben,  1777 

Cebus  curtus  Bangs,  1905b 
Bull.  Mus.  Comp.  ZooL,  46:  91,  June. 
=  Cebus  capucinus  curtus  Bangs,  1905. 
See  Cabrera  (1917:  240). 

Holotype.  MCZ  10824.  Skin  and  skull.  Adult  male. 
Locality.    Colombia:    (Cauca),   Gorgona   Island.   2 
July  1904. 

Collector.  W.  W.  Brown,  Jr.  Original  number  27. 
Condition.  Skin  and  skull  complete. 
Type   Series.    1   paratype;    MCZ   10825,   skin   and 
skull,  adult  female. 

Comments.  C.  c.  curtus  was  retained  as  a  valid  sub- 
species by  Cabrera  (1958:  169)  and  Napier  (1976: 
36). 

Family  HYLOBATIDAE  Gray,  1870 

Genus  HYLOBATES  llliger,  1811 

Hylobates  lar  carpenteri  Groves,  1968 
Proc.  Biol.  Soc.  Washington,  81:  625,  30 
December. 

Holotype.    MCZ    41430.    Skin    and    skeleton.    Adult 
male. 

Locality.  Thailand:  Chiengmai  District,  Mount 
Angka  (  =  Doi  Inthanon),  3,400  ft  (1,037  m).  14 
March  1937. 

Collector.  H.  J.  Coolidge,  Jr.,  Asiatic  Primate  Ex- 
pedition. Original  numbers  21  and  185. 
Condition.  Skin,  skull,  and  skeleton  complete. 
Type  Series.  Groves  examined  a  series  of  144  skins, 
skulls,  and  skeletons  of  this  subspecies  that  were 
collected  by  the  Harvard  Asiatic  Primate  Expedi- 
tion in  1937  from  Chiengmai  District,  Thailand, 
"mostly  in  the  Museum  of  Comparative  Zoology, 
Hai'vard,  but  a  few  of  the  osteological  specimens 
in  the  Anthropology  Department,  University  of 
California  at  Berkeley."  He  examined  also  18  spec- 
imens in  the  USNM  and  1  in  the  AMNH,  all  rep- 
resented by  skin  and  skull. 

Comments.  Retained  as  a  valid  subspecies  by  Jen- 
kins (1990:  15)  and  Geissman  (1995:  474). 

Family  HOMINIDAE  Gray,  1825 

Genus  GORILLA  I.  Geoff roy,  1853 

Troglodytes  gorilla  Savage  and  Wyman, 

1847 

Boston  J.  Nat.  Hist.,  5:  417,  December. 

=  Gorilla  gorilla  gorilla  (Savage  and 

Wyman,  1847).  See  Rothschild  (1923: 

176). 

Lectotype.  MCZ  9587.  Skull  and  postcranial  skeleton. 
Adult  male. 


Locality.  Gabon:  Gabon  Estuaiy,  Mpongwe  coun- 
tiy.  1847. 

Collector  T.  S.  Savage.  Original  Jiumber  28  (of  J. 
Wyman). 

Condition.  Postcranial  skeleton  partial  (includes 
pelvis,  sacrum,  scapulae,  humeri,  radii,  left  ulna, 
femora,  left  tibia,  and  7  vertebrae:  2  cervical,  3  dor- 
sal, 2  lumbar).  Head  of  right  humerus  and  left  fe- 
mur bisected. 

Type  Series.  The  original  description  mentions 
"four  skulls,  two  males  and  two  females,  one  of 
each  in  a  perfect  condition,  and  all  of  them  adult; 
a  male  and  female  peKis,  the  long  bones  of  the 
upper  and  lower  extremities,  and  a  few  vertebrae 
and  ribs."  However,  measurements  and  illustrations 
of  only  two  of  these  specimens,  an  adult  male  and 
female,  each  represented  by  a  skull  and  partial 
postcranial  skeleton,  are  provided.  These  two,  pre- 
sumably the  specimens  originally  described  as  be- 
ing "in  a  perfect  condition,"  are  the  only  specimens 
of  the  original  four  to  be  noted  in  Wymans  per- 
sonal notebook  of  osteology  (unpublished,  now  in 
tlie  libraiy  of  the  Boston  Museum  of  Science)  and 
die  only  original  specimens  of  which  there  is  any 
record  at  all.  These  two  specimens  were  trans- 
ferred from  the  Boston  Society  of  Natural  History 
to  the  MCZ  in  1915-16.  They  bear  MCZ  numbers 
9587  (male)  and  9311  (female).  The  skull  of  the 
female  was  sawed  in  half  (hemisected),  probably  by 
Wyman.  The  adult  male  specimen,  MCZ  9587, 
possesses  an  intact  skull.  MCZ  9587  is  the  only  in- 
tact specimen  of  the  syntype  series  known  to  exist, 
and  its  measurements  and  an  illustration  of  its  skull 
are  included  in  the  original  publication.  Addition- 
ally, it  is  the  only  syntype  whose  measurements  are 
included  in  Coolidge  s  rex-ision  of  the  genus  Gorilla 
(1929:  325),  although  it  is  erroneously  listed  in  that 
work  as  MCZ  9586.  For  these  reasons,  MCZ  9587 
is  hereby  designated  as  lectotype  of  Troglodytes  go- 
rilla Savage  and  Wyman,  1847;  this  should  ensure 
taxonomic  consistency  between  past  and  future 
treatments  of  this  name. 

Comments.  Type  species  of  the  genus  Gorilla  I. 
Geoffroy,  1853.  The  original  description  is  occa- 
sionally attributed  in  error  to  Wyman  (1847).  G. 
gorilla  was  considered  a  valid  species  by  Groves 
a993:  276)  and  Nowak  (1999:  618). 

Order  LIPOTYPHLA  Haeckel,  1866 

Family  NESOPHONTIDAE  Anthony,  1916 

Genus  /VESOPHOA/7ES  Anthony,  1916 

Nesophontes  micrus  G.  M.  Allen,  1917a 
Bull.  Mus.  Comp.  ZooL,  61:  5,  Januaiy. 

Holotype.  VP  9600.  Right  ramus. 

Locality.  Cuba:  Matanzas  Province,  cave  in  Sierra 
de  Hato  Neuvo. 
Collector  C.  de  la  Torre. 


Type  Specimens  of  Recent  Mammals  •  Helgen  and  McFadden        137 


Condition.    Condition    as    originally    described — 
"posterior  half  of  the  right  ramus,  containing  a  part 
o£ pm^,  m„  m.2,  and  the  roots  of  111.." 
Type  Series.  Holotype  only. 

Comments.  Considered  a  valid  species  by  Hutterer 
(1993:  70)  and  Nowak  (1999:  201);  almost  certainly 
extinct.  This  specimen  is  stored  in  the  Vertebrate 
Paleontology  Department  of  the  MCZ. 

Family  SOLENODONTIDAE  Gill,  1872 

Genus  SOL EA/ODOA/ Brandt,  1833 

Antillogale  marcanoi  Patterson,  1962 

Breviora,  Mus.  Comp.  ZooL,  165:  3,  22 

August. 

=  Solenodon  marcano/ (Patterson,  1962). 

See  Varona  (1974:  8). 

Holotype.  VP  7261.  Right  lower  ramus. 

Locality.  Dominican  Republic:  San  Rafael  Prov- 
ince (  =  Elias  Pina),  Hondo  Valle  Municipality,  un- 
named cave  2  km  southeast  of  Rancho  La  Guardia. 
Summer  1958. 

Collector.  C.  E.  Ray  and  A.  S.  Rand. 
Condition.  Condition  as  originally  described — "in- 
complete right  ramus  of  mandible  with  P3— M,  and 
alveoli  of  other  teeth." 

Type  Series.  5  paratypes;  7262,  left  ramus,  7263, 
right  humerus,  7264,  left  humerus,  7265,  right 
ulna,  7266,  left  ramus,  juvenile;  all  partial. 
Comments.  A.  marcanoi  is  the  type  species  of  the 
genus  Antillogale  Patterson,  1962.  Antillogale  was 
first  synonymized  with  Solenodon  by  Van  Valen 
(1967:  255).  Considered  a  valid  species  by  Hutterer 
(1993:  69)  and  Nowak  (1999:  199).  S.  marcanoi  is 
probably  extinct.  This  specimen  is  stored  in  the 
Vertebrate  Paleontology  Department  of  the  MCZ. 

Solenodon  poeyanus  Barbour,  1944 
Proc.  New  England  Zool.  Club,  23:  6, 
March  7. 

=  Solenodon  cubanus  poeyanus  Barbour, 
1944.  See  Aguayo  (1950:  131). 

Holotype.  MCZ  6597.  Mounted  skin  and  skull. 

Locality.  Cuba:  Oriente  (  =  Holguin),  near  Nipi  Bay 
(  =  Nipe  Bay). 

Collector  Bought  by  A.  Agassiz  from  H.  A.  Ward, 
1891. 

Condition.  Skin  complete.  Skull  partial  (basioccip- 
ital  plate  and  tympanic  bullae  missing,  parietals 
damaged).  Mandible  disarticulated. 
Type  Series.  In  the  original  description,  Barbour 
includes  a  photograph  of  a  "living  example  of  So- 
lenodon poeyanus  now  in  Zoological  Garden,  Ha- 
vana, Cuba,  from  vicinity  of  Baracoa." 
Comments.  The  number  of  this  specimen  is  6597, 
not  6957  as  stated  in  the  original  description.  S.  c. 


poeyanus  was  retained  as  a  valid  subspecies  by  Hall 

(1981:  22). 

Family  SORICIDAE  Fischer  von 
Waldheim,  1817 

Genus  BLARINA  Gray,  1838 

Blarina  brevicauda  aloga  Bangs,  1902a 
Proc.  New  England  Zool.  Club,  3:  76,  31 
March. 

Holotype.  B9727.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Massachusetts,  Dukes 
County,  Marthas  Vineyard,  West  Tisbury.  25  June 
1899. 

Collector  O.  Bangs.  Original  number  2. 
Condition.  Skin  and  skrdl  complete.  Mandible  dis- 
articulated. 

Type  Series.  9  paratypes,  B9725,  B9726,  B9728- 
B9734,  all  represented  by  skin  and  skull,  3  females 
and  6  feinales. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  54)  and  George  et  al.  (1986:  1). 

Blarina  brevicauda  compacta  Bangs, 

1902a 

Proc.  New  England  Zool.  Club,  3:  77,  31 

March. 

Holotype.  B9705.  Skin  and  skull.  Adult  male. 

Locality.    (United    States):    Massachusetts,    (Nan- 
tucket County),  Nantucket  (Island).  10  July  1899. 
Collector  O.  Bangs.  Original  number  3. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  6  paratypes;  B9701-B9704,  B9706, 
B9708;  all  represented  by  skin  and  skull,  4  females 
and  2  males. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  56)  and  George  et  al.  (1986:  1). 

Genus  CROCIDURAVJagner,  1832 

Crocidura  bicolor  tephragaster  Setzer, 

1956 

Proc.  U.S.  Nat.  Mus.,  106:  458,  28 

November. 

=  Crocidura  fuscomurina  (Heuglin,  1865). 

See  Hutterer  (1983:  223). 

Holotype.  MCZ  44773.  Skin  and  skull.  Adult  male. 
Locality.  Anglo-Egyptian  Sudan  (  =  Sudan):  (East- 
ern) Equatoria,  Torit.  25  April  1950. 
Collector.  J.  S.  Owen.  Original  number  1158. 
Condition.  Sldn  and  skull  complete. 
Type  Series.  Setzer  mentions  that  he  examined  18 
specimens  of  tephragaster,  8  of  which  are  in  the 
MCZ. 


138         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Crocidura  hildegardeae  phaios  Setzer, 

1956 

Proc.  U.S.  Nat.  Mus.,  106:  460,  28 

November. 

Holotype.  MCZ  45S55.  Skin  and  skull.  Adult  female. 
Locality.  Anglo-Egyptian  Sudan  (  =  Sudan):  Equa- 
toria,  Imatong  Mts,  Gilo,  6,500  ft  (1,983  m).  12 
June  1950. 

Collector.  J.  S.  Owen.  Original  number  1266. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  3  paratypes;  1  from  the  MCZ;  MCZ 
45856,  skin  and  skull,  male. 

Comments.  Retained  as  a  valid  subspecies,  C  gra- 
cilipes  phaios,  by  Heim  de  Balsac  and  Meester 
(1977:  16);  included  in  Crocidura  hildegardeae  by 
Hutterer  (1993:  87). 

Genus  CRYPTOTIS  Pomel,  1848 

Cryptotis  avia  G.  M.  Allen,  1923a 

Proc.  New  England  Zool.  Club,  S:  37,  12 

February. 

=  Cryptotis  thomasi  {Mernam,  1897).  See 

Woodman  (1996:  414). 

Holotype.  MCZ  20091.  Skin  and  skull.  Adult. 

Locality.  Colombia:  (probably  Cundinamarca),  El 
Verjon  (see  coments).  October  1922. 
Collector  N.  Maria. 

Condition.    Sldn  complete.   Skull  partial  (most  of 
skull  from  frontals  to  occiput  missing).  Left  ramus 
of  mandible  missing. 
Type  Series.  Holotype  only. 

Comments.  Considered  a  valid  species  by  Hutterer 
(1993:  108)  and  Nowak  (1999:  209)  but  synony- 
mized  as  noted  above.  The  locality  "El  Verjon"  is 
not  shown  on  any  maps  available  to  us  but  is  pos- 
sibly equivalent  to  "Paramo  Cruz  Verde"  (Paynter 
1997:  463). 

Genus  MYOSOREX  Gray,  1838 

Crocidura  maurisca  geata  G.  M.  Allen  and 

Loveridge,  1927 

Proc.  Boston  Soc.  Nat.  Hist.,  38:  417,  23 

December. 

=  Myosorex  geata  (G.  M.  Allen  and 

Loveridge,  1927).  See  Heim  de  Balsac 

(1967:  610). 

Holotype.  MCZ  22447.  Skin  and  skull.  Adult  male. 
Locality.  Tanganyika  Territoiy  (=  Tanzania):  Mor- 
ogoro,    Uluguru    Mountains,    Nyingwa,    7,500    ft 
(2,288  m).  19  October  1926. 
Collector  A.  Loveridge. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 


Type   Series.    1   paratype;    MCZ  22448,   skin   and 
skull,  adult  female. 

Comments.  M.  geata  was  considered  a  valid  species 
by  Hutterer  (1993;  99)  and  Nowak  (1999:  217). 

Genus  SOREX  Linnaeus,  1758 

Neosorex  palustris  acadicus  G.  M.  Allen, 

1915a 

Proc.  Biol.  Soc.  Washington,  28:  15,  12 

February. 

Name  preoccupied  by  Sorex  acadicus 
Gilpin,  1867. 

Sorex  palustris  gloveralleni  Jackson,  1 926 
J.  Mammal.,  7:  57,  15  February. 
(Replacement  name  for  Neosorex 
palustris  acadicus  G.  M.  Allen,  1915) 

Holotype.  B2046.  Skin  and  skull.  Adult  female. 

Locality.    (Canada):   Nova   Scotia,   Digby.   26  July 
1894. 

Collector  O.  Bangs.  Original  number  3. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  1  paratype;  B2053,  exchanged  to  the 
USNM  in  1922. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  41)  and  Beneski  and  Stinson  (1987:  1). 

Sorex  araneus  ultimus  G.  M.  Allen,  1914b 

Proc.  New  England  Zool.  Club,  5:  51,  9 

April. 

=  Sorex  tundrensis  Merriam,  1900.  See 

Hutterer  (1993:  121). 

Holotype.  MCZ  15000.  Skin  and  skull.  Adult  male. 
Locality.  (Russian  Federation):  northeastern  Sibe- 
ria, Nijni  Kolymsk  (  =  Nizhnekolymsk),  near  mouth 
of  Kolyma  River.  6  November  1911. 
Collector.  J.  Koren.  Original  number  136. 
Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  There  is  a  series  of  paratypes,  most  of 
which  are  housed  in  the  MCZ. 

Sorex  macropygmaeus  koreni  G.  M.  Allen, 

1914b 

Proc.  New  England  Zool.  Club,  5:  56,  9 

April. 

=  Sorex  caecutiens  koreni  G.  M.  Allen, 

1914.  See  Yudin  (1989:  281). 

Holotype.  MCZ  15085.  Skin  and  skiill.  Adult  female. 
Locality.  (Russian  Federation):  northeastern  Sibe- 
ria, Nijni  Kolymsk  (  =  Nizhnekolymsk),  near  mouth 
of  Kolyma  River.  19  October  1911. 
Collector  J.  Koren.  Original  number  .50. 


Type  Specimens  of  Recent  Mammals  •  Helaen  and  McFadden 


139 


Tijpe  Series.  5  paratypes;  MCZ  15003-15007,  all 
represented  by  sldn  and  skull;  4  males,  1  female. 
MCZ  15004,  a  male,  was  exchanged  to  the  FMNH 
in  1931. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Sorex  macrurus  Batchelder,  1896 

Proc.  Biol.  Soc.  Washington,  10:  133,  8 

December. 

Name  preoccupied  by  Sorex  macrourus 

Lehmann,  1822. 

Sorex  dispar  Batchelder,  191 1 
Proc.  Biol.  Soc.  Washington,  24:  97,  15 
May.  (Replacement  name  for  Sorex 
macrurus  Batchelder,  1896) 

Holotype.  MCZ  41744.  Skin  and  skaill.  Adult  male. 
Locality.  (United  States):  New  York,  Essex  County, 
Keene  Heights,  Beede's  (see  comments).  9  Sep- 
tember 1895. 

Collector.  C.  F.  Batchelder.  Original  number  1384. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  1  paratype;  MCZ  41745,  skin  and 
skull,  adult  male. 

Comments.  S.  dispar  was  considered  a  valid  species 
by  Hutterer  (1993:  114)  and  Nowak  (1999:  205). 
The  type  loctility  was  redescribed  by  Martin  (1966: 
131)  as  "0.6  mile  south  and  0.5  mile  east  of  Saint 
Huberts,  Essex  County,  New  York,  lat.  44°09',  long. 
73°46'." 

Sorex  personatus  miscix  Bangs,  1899d 

Proc.  New  England  Zool.  Club,  1:  15,  28 

Februaiy. 

=  Sorex  cinereus  miscix  Bangs,  1899.  See 

Jackson  (1925:  56). 

Holotype.  B8651.  Skin  and  skull.  Adult  male. 

Locality.  (Canada):  Labrador,  Black  Bay.  10  Octo- 
ber 1898. 

Collector  E.  Doane.  Original  number  1. 
Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type   Series.    Bangs  based  his   description  on   39 
specimens;    corresponding   to    B7931-B7950    and 
B8651-B8669;  all  represented  by  skin  and  skull. 
Comments.  S.  c.  miscix  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  29). 

Sorex  sanguinidens  G.  M.  Allen,  1914b 

Proc.  New  England  Zool.  Club,  5:  54,  9 

April. 

=  Sorex  daptiaenodon  sanguinidens  G.  M. 

Allen,  1914.  See  Yudin  (1989:  198). 

Holotype.  MCZ  15012.  Skin  and  skull.  Adult  female. 
Locality.  (Russian  Federation):  northeastern  Sibe- 


ria, Nijni  Kolymsk  (  =  Nizhnekolymsk),  near  mouth 

Kolyma  River.  11  December  1911. 

Collector  J.  Koren.  Original  number  221. 

Condition.  Tail  separate  from  sldn.  Skull  complete. 

Mandible  disarticulated. 

Type  Series.  There  is  a  large  series  of  paratypes, 

most  of  which  are  still  in  the  MCZ. 

Sorex  virG.  M.  Allen,  1914b 

Proc.  New  England  Zool.  Club,  5:  52,  9 

April. 

=  Sorex  roboratus  Hollister,  1913.  See 

Hoffman  (1985:  17). 

Holotype.  MCZ  15068.  Skin  and  skull.  Adult  female. 
Locality.  (Russian  Federation):  northeastern  Sibe- 
ria, Nijni  Kolymsk  (  =  Nizhnekolymsk),  near  mouth 
of  Kolyma  River.  19  December  1911. 
Collector  J.  Koren.  Original  number  230. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  There  is  a  large  series  of  paratypes, 
most  of  which  are  still  in  the  MCZ. 

Genus  SL/A/CL/S  Ehrenberg,  1832 

Suncus  aterMedway,  1965 

Mammals  of  Borneo,  J.  Malay.  Branch  R. 

Asiat.  Soc,  36:  38,  December. 

Holotype.    MCZ  36574.   Skin,  skull,  and  postcranial 
skeleton.  Adult  female. 

Locality.  Malaysia:  northern  Borneo,  Sabah, 
Mount  Kinabalu,  Lumu  Lumu,  5,500  ft  (1,678  m). 
7  July  1937. 

Collector  J.  A.  Griswold,  Jr.  Original  number  462. 
Condition.  Skin  complete.  Skull  partial  (tympanic 
bullae  missing),  and  mandible  disarticulated.  Post- 
cranial  skeleton  complete. 
Type  Series.  Holotype  only. 

Comments.  Considered  a  valid  species  by  Hutterer 
(1993:  101)  and  Nowak  (1999:  223). 

Suncus  varilla  minor  G.  M.  Allen  and 

Loveridge,  1933 

Bull.  Mus.  Comp.  Zool,  75:  57, 

Februaiy. 

Holotype.  MCZ  26754.  Skin  and  skaill.  Adult  female. 
Locality.  Tanganyika  Territory  (=Tanzania):  Urun- 
gu,  Kitungulu,  4,500  ft  (1,373  m).  14  May  1930. 
Collector.  A.  Loveridge. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Holotyj^e  only. 

Comments.  Retained  as  a  valid  subspecies  by  Heim 
de  Balsac  and  Meester  (1977:  6). 


140         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Family  TALPIDAE  Fischer  de  Waldheim, 
1817 

Genus  NEUROTRICHUS  Gm\her,  1880 

Neurotrichus  gibbsi  hyacinthinus  Bangs, 

1897d 

Amer.  Nat.,  31:  240,  1  March. 

Holotype.  B1240.  Sldn  and  skull.  Adult  female. 

Locality.  U.S.A.  (United  States):  California,  Marin 
County,  Nicasio.  10  March  1894. 
Collector.  C.  A.  Allen.  Original  number  694. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  1  paratype;  B1241,  skin  and  skull, 
adult  male. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  67)  and  Carraway  and  Verts  (1991:  1). 

Genus  SCALOPUS  Desmarest,  1 804 

Scalops  anastasae  Bangs,  1898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  212,  15 

March. 

=  Scalopus  aquaticus  anastasae  (Bangs, 

1898).  See  Jackson  (1915:  39). 

Holotype.  B7192.  Skin  and  skull.  Adult  male. 

Locality.  U.S.A.  (United  States):  Florida,  St.  Johns 
County,  Anastasia  Island,  Point  Romo.  16  Febioiary 
1897. 

Collector  O.  Bangs.  Original  number  10. 
Condition.  Skin  and  skaiU  complete.  Mandible  dis- 
articulated. 

Tijpe  Series.  4  paratypes;  B719.3-B7196;  all  repre- 
sented by  sldn  and  skull,  2  females  and  2  males. 
Comments.  S.  a.  anastasae  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  72). 

Scalops  texanus  aereus  Bangs,  1896li 

Proc.  Biol.  Soc.  Washington,  10:  138,  28 

December. 

=  Scalopus  aquaticus  aereus  (Bangs, 

1896).  See  Miller  (1912:  8). 

Holotype.  B547.5.  Skin  and  skull.  Adult  female. 

Locality.  (United  States):  Indian  Territoiy 
(  =  Oklalioma)  (Adair  County),  Stilwell.  13  August 
1896. 

Collector.  T.  Surber.  Original  number  64. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Holotype  only. 

Comments.  S.  a.  aereus  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  72). 


Genus  SCAPANUS  Pome\,  1848 

Scapanus  californicus  minusculus  Bangs, 

1899J 

Proc.  New  England  Zool.  Club,  1:  70,  31 

Jnly. 

=  Scapanus  latimanus  minusculus  Bangs, 

1899.  See  Grinnell  and  Swarth  (1912: 

133). 

Holotype.  B9189.  Skin  and  skull.  Adult  female. 
Locality.    (United   States):   California,   El   Dorado 
County,  Fyffe.  10  June  1897. 

Collector  W.  W.  Price  and  E.  M.  Nutting.  Original 
number  15. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Holotype  only. 

Comments.  S.  I.  minusculus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  70). 

Order  CHIROPTERA  Blumenbach,  1779 

Family  PTEROPODIDAE  Gray,  1821 

Genus  AETHALOPS  Thomas,  1923 

Aethalops  aequalis  G.  M.  Allen,  1938b 
J.  Mammal.,  19:  497,  14  November. 
= Aethalops  alecto  aequalis  G.  M.  Allen, 
1938.  See  Hill  (1961:  639). 

Holotype.    MCZ  36582.   Skin,  skull,  and  postcranial 
skeleton.  Adult  female. 

Locality.  (Malaysia):  British  North  Borneo  (  =  Sa- 
bali).  Mount  Kinabalu,  Luma  Luma,  5,500  ft  (1,678 
m).  12  July  1937. 

Collector.  J.  A.  Griswold,  Jr.,  Asiatic  Primate  E.x;- 
pedition.  Original  number  510. 
Condition.    Sldn,    skull,    and   postcranial   skeleton 
complete. 

Type  Senes.  3  paratypes;  MCZ  36583,  sldn,  skull 
and  skeleton,  female;  MCZ  36584,  skin  and  skull, 
female;  MCZ  36586,  skin  and  skull,  female. 
Comments.  A.  a.  aequalis  was  retained  as  a  valid 
subspecies  by  Koopman  (1994:  35). 

Genus  EONYCTERIS  Dobson,  1873 

Eonycteris  spelaea  glandifera  Lawrence, 

1939 

7/7  Barbour,  Lawrence,  and  Peters,  Bull. 

Mus.  Comp.  Zool.,  86:  38,  November. 

Holotype.  MCZ  35159.  Sldn  and  skull.  Adult  male. 
Locality.  Philippines:  Luzon,  (Nueva  Ecija),  Rizal, 
Montalban  caves  near  Manila.  27  Febiniary  1937. 
Collector.  B.  Lawrence.  Original  number  253. 
Condition.  Skin  and  skull  complete. 
Type  Series.  In  the  original  description,  Lawrence 


Type  Specimens  of  Recent  Mammals  •  Helaen  and  McFadden 


141 


implies  the  existence  of  several  specimens  in  ad- 
dition to  the  holotype;  she  collected  18  specimens 
of  glandifera  during  her  1936—37  expedition  to  the 
Philippines,  all  of  which  are  in  the  MCZ. 
Comments.  Retained  as  a  valid  subspecies  by  Ma- 
haradatunkamsi  and  Kitchener  (1997:  59). 

Genus  HAPLONYCTERIS  Lawrence, 
1939 

Haplonycteris  fischeri  Lawrence,  1939 
In  Barbour,  Lawrence,  and  Peters,  BulL 
Mus.  Comp.  ZooL,  86:  33,  November. 

Holotype.  MCZ  35258.  Skin  and  skull.  Adult  male. 
Locality.   Philippines:   Mindoro  (Oriental),  Mouirt 
Halcon,  Bignay.  26  April  1937. 
Collector  F.  S.  Rivera.  Original  number  BL  502. 
Condition.  Skin  and  skull  complete. 
Type  Series.  Holotype  only. 

Comments.  Type  species  of  the  genus  Haplonyc- 
teris Lawrence,  1939.  Considered  a  valid  species 
by  Koopman  (1993:  142)  and  Nowak  (1999:  292). 

Genus  PTEROPUS  Erxleben,  Mil 

Pteropus  anetianus  aorensis  Lawrence, 

1945 

Proc.  New  England  ZooL  Club,  23:  66, 

26  March. 

Holotype.  MCZ  42183.  Skin  and  skull.  Adult  male. 
Locality.  (Vanuatu):  New  Hebrides,  off  southwest 
corner  of  Espiritu   Santo   Island,   Aore   Island.   8 
April  1944. 

Collector  O.  L.  Austin,  Jr.  Original  number  5. 
Condition.  Skin  complete.  Skull  partial  (basioccip- 
ital  missing),  right  ramus  of  mandible  broken. 
Type   Series.    1   paratype;    MCZ  42182,   skin   and 
skull,  adult  male. 

Comments.  Retained  as  a  valid  subspecies  by 
Koopman  (1994:  25)  and  Flanneiy  (1995b). 

Pteropus  arielG.  M.  Allen,  1908 
Bull.  Mus.  Comp.  ZooL,  52:  28,  July. 
=  Pteropus  giganteus  ariel  G.  M.  Allen, 
1908.  See  Hill  (1958:  5). 

Holotype.  MCZ  10565.  Skin  and  skull.  Adult  male. 
Locality.  Maldive  Islands:  Male  Atoll.  24  Decem- 
ber 1901. 

Collector.  H.  B.  Bigelow,  A.  Agassiz  E.xpedition. 
Condition.  Right  wing  of  sldn  damaged.  Skull  in- 
tact, with  2  small  holes  in  braincase. 
Type   Series.    1   paratype;    MCZ   10566,   skin   and 
skull,  juvenile  female. 

Comments.  P.  g.  ariel  was  retained  as  a  valid  sub- 
species by  Koopman  (1994:  26). 


Pteropus  austini  Lawrence,  1945 

Proc.  New  England  Zool.  Club,  23:  59, 

26  March. 

=  Pteropus  woodfordi  Thomas,  1888.  See 

Sanborn  and  Beecher  (1947:  389). 

Holotype.   MCZ  42166.  Skin  and  skull.  Subadult  fe- 
male. 

Locality.  Solomon  Islands:  Florida  Island  (Nggela 
Group).  20  Februaiy  1944. 
Collector  O.  L.  Austin,  Jr.  Original  number  2. 
Condition.  Skin  and  skull  complete. 
Type   Series.    1   paratype;    MCZ  42167,   skin   and 
skull,  subadult  male. 

Comments.  Lawrence  referred  to  the  holotype  as 
an  adult  in  the  original  description,  but  Sanborn 
and  Beecher  (1947:  389)  recognized  it  as  a  sub- 
adult. 

Pteropus  rayneri  monoensis  Lawrence, 

1945 

Proc.  New  England  Zool.  Club.,  23:  63, 

26  March. 

Holotype.  MCZ  42191.  Skin  and  skull.  Aduh  male. 
Locality.  Solomon  Islands:  Treasuiy  (Mono)  Island. 
11  October  1944. 

Collector  O.  L.  Austin,  Jr.  Original  number  27. 
Condition.  Skin  and  skull  complete. 
Type   Series.   2  paratypes;   MCZ  42192,  skin  and 
skull,  adult  male;  MCZ  42193,  skin  and  skull,  adult 
male. 

Comments.  Retained  as  a  valid  subspecies  by 
Koopman  (1994:  24)  and  Flannery  (1995b:  285). 

Genus  ROUSETTUS  Gray,  1821 

Rousettus  madagascariensis  G. 

Grandidier,  1930b 

Bull.  Acad.  Malgache,  n.  sen,  11:  91  (for 

1928). 

Holotype.    MCZ  45432.   Alcoholic   and  skull.   Adult 
male. 

Locality.  Malagasy  Republic  (Madagascar):  (Anta- 
nanarivo), between  Tananarive  (  =  Antananarivo) 
and  Andevoranto,  Grand  forest  de  Est,  near  Be- 
forona. 

Collector    Received   by   G.    Grandidier  from   the 
Academic  Malgache,  1917. 
Conditio72.  Alcoholic,  skull  complete. 
Type  Series.  Holotype  only. 

Comments.  Considered  a  valid  species  by  Koop- 
man (1993:  153)  and  Nowak  (1999:  261).  In  his 
review  of  known  material  of  jR.  madagascariensis, 
Bergmans  (1977:  67)  commented  in  error  that  the 
holotype  was  in  the  Academic  Malgache,  Antana- 
narivo, Madagascar. 


142         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Family  EMBALLONURIDAE  Gervais,  1855 

Genus  PEROPTERYX  Peters,  1867 

Peropteryx  canina  phaea  G.  M.  Allen, 

1911a 

Bull.  Mus.  Comp.  ZooL,  54:  222,  July. 

=  Peropteryx  macrotis  phaea  G.  M.  Allen, 

1911.  See  Sanborn  (1937:  342). 

Holotijpe.  MCZ  8101.  Skin  and  skull.  Adult  female. 
Locality.  Grenada:  Point  Saline(s).  29  August  1910. 
Collector.  G.  M.  Allen.  Original  number  15. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  P.  m.  phaea  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  82)  and  Koopman  (1994: 
47). 

Genus  RHYNCHONYCTERIS  Peters, 
1867 

Rhynchiscus  naso  priscus  G.  M.  Allen, 

1914c 

Proc.  Biol.  Soc.  Washins^ton,  27:  109,  10 

July. 

=  Rhynchonycteris  naso  (Wied-Neuwied, 

1820).  See  Sanborn  (1937:  326). 

Holotijpe.  MCZ  13208.  Skin  and  skull.  Adult. 

Locality.  Me.\ico:  Quintana  Roo,  Xcopen.  18  Feb- 
ruary 1912. 

Collector.  J.  L.  Peters.  Original  number  13. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  3  paratypes;  MCZ  13209,  skin  and 
skuU,  adult  male;  MCZ  14637,  alcoholic,  adult 
male;  MCZ  14638,  alcoholic,  adult  female. 

Family  NYCTERIDAE  Van  der  Hoeven, 
1855 

Genus  NYCTERIS  G.  Cuvier  and  E. 
Geoffroy,  1795 

Nycteris  madagascariensis  G.  Grandidier, 

1937 

Bull.  Mus.  Nat.  Hist.  Paris,  9:  353,  25 

November. 

Holotype.  MCZ  45433.  Body  in  alcohol  skull  extract- 
ed. 

Locality.  Madagascar:  (Antsiranana),  Diego-Suarez 
(  =  Antsiranana),  Valley  of  the  Rodo,  north  of  Pir- 
kana  near  the  Ankarana,  12°5'-13°0'S,  49°5'E.  June 
1910. 
Collector.  Grandidier  collection. 


Condition.  Alcoholic,  skull  complete. 

Type  Series.   1  paratype,  MCZ  45434,  in  alcohol, 

skull  extracted,  female. 

Comments.  N.  madagascariensis  was  included  in  N. 

macrotis  by  Koopman  (1993:  162)  but  retained  as 

a  valid  species  by  Peterson  et  al.  (1995;  6.3). 

Nycteris  nana  tristis  G.  W\.  Allen  and 
Lawrence,  1936 

Bull.  Mus.  Conip.  Zool.,  79:  47,  January. 
^Nycteris  nana  (Andersen,  1912).  See 
Hayman  and  Hill  (1971:  19). 

Holotype.  MCZ  31156.  Skin  and  skull.  Adult  female. 
Locality.  Kenya:  (W.  Nyanza),  Kakamega  District, 
Kaimosi.  13  Februaiy  1934. 
Collector  A.  Loveridge. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 
Type  Series.  Holotype  only. 

Family  RHINOLOPHIDAE  Gray,  1825 

Genus  HIPPOSIDEROSGmy,  1831 

Hipposideros  curtus  G.  M.  Allen,  1921 
Rev.  Zool.  Africaine,  9:  194,  December. 

Holotype.  MCZ  19305.  Body  in  alcohol.  Female. 
Locality.  Cameroons  (  =  Cameroon):  (Littoral),  Sak- 
bayeme.  1920. 
Collector  G.  Schwab. 

Condition.    Alcoholic.    Skull    extracted   but    not 
cleaned  (still  in  alcohol). 
Type  Series.  Holotype  only. 

Comments.  H.  curtus  was  considered  a  valid  spe- 
cies by  Koopman  (1993:  172)  and  Nowak  (1999: 
333).  ' 

Hipposideros  erigens  Lawrence,  1939 
In  Barbour,  Lawrence,  and  Peters,  Bull. 
Mus.  Comp.  Zool.,  86:  56,  November. 
=  l-lipposideros  bicolor  erigens  Lawrence, 
1939.  See  Hill  (1963:  28). 

Holotype.  MCZ  35197.  Skin  and  skull.  Adult  male. 
Locality.  Philippines:  Mindoro,  (Oriental)  Tabucala 
cave  near  Calapan,  northern  base  of  Mount  Hal- 
con.  7  March  1937. 

Collector  B.  Lawrence.  Original  number  307. 
Condition.  Skin  complete.  Skull  partial  (occiput 
missing),  and  mandible  disarticulated. 
Type  Series.  3  paratypes;  MCZ  35195,  skin  and 
skull,  adult  female;  MCZ  35196,  sldn  and  skull, 
adult  male;  MCZ  35198,  skin  and  skull,  adult  fe- 
male. 

Comments.  H.  b.  erigens  was  retained  as  a  valid 
subspecies  by  Koopman  (1994;  61). 


Type  Specimens  of  Recent  Mammals  •  Helaen  and  McFadden 


143 


Hipposideros  turpis  Bangs,  1901a 
Amer.  Nat.,  35:  561,  31  July. 
=  Hipposideros  turpis  turpis  Bangs,  1901. 
See  Hill  (1963:  94). 

Holotijpe.  MCZ  10003.  Skin  and  skull.  Adult  female. 
Locoliti/.  (Japan):  Ryaikyu  Islands,  southern  group 
of  Liu  kin  Islands,  Ishigaki  Island.  10  May  1899. 
Collector.  I.  Zensaku. 

Condition.  Sldn  complete.  Skull  partial  (occipital 
region  missing). 

Type  Series.  2  paratypes;  MCZ  10002,  skin  and 
skull,  adult  female;  MCZ  10004,  skin  and  skvill, 
adult  male. 

Comments.  H.  tuiyis  was  considered  a  valid  species 
by  Koopman  (1993:  175)  and  Nowak  (1999:  334). 

Genus  RHINOLOPHUS  Lacepe6e,  1799 

Riiinoloplius  megaphyllus  igniter  G.  M. 

Allen,  1933 

J.  Mammal.,  14:  149,  15  May. 

=  Rtiinolophus  megapfiyilus  megapiiyllus 

Gray,  1834.  See  Koopman  (1984:  9). 

Holotijpe.  MCZ  29078.  Skin  and  skull.  Adult  male. 
Locality.  Australia:  Queensland,  Cape  York,  Coen. 
12  June  1932. 

Collector  P.  J.  Darlington,  Jr.,  Hai-vard  Australian 
Expedition.  Original  number  185. 
Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  1  paratyj^jc;  MCZ  29079,  skin  and 
skull,  adult  male. 

Rtiinolophus  ptiilippinensis  alleni 
Lawrence,  1939 

In  Barbour,  Lawrence,  and  Peters,  Bull. 
Mus.  Comp.  Zool.,  86:  46,  November. 

Holotype.  MCZ  35097.  Sldn  and  skull.  Adult  female. 
Locality.  Philippines:  Mindoro,  (Oriental)  Tabucala 
cave  near  Calapan,  northern  base  of  Mount  Hal- 
con.  7  March  1937. 

Collector.  B.  Lawrence.  Original  number  302. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  2  pai'atypes;  MCZ  35098,  skin  and 
sk-ull,  adult  female;  MCZ  35099,  skin  and  skull, 
adult  female. 

Comments.  Retained  as  a  valid  subspecies  by 
Koopman  (1994:  57). 

Genus  TRIAENOPS  Dobson,  1871 

Triaenops  aurita  G.  Grandidier,  1912 
Bull.  Mus.  Hist.  Nat.  Paris,  18:  8,  25 
Januaiy. 

Holotype.  MCZ  45080.  Mummy. 

Locality.  Malagasy  Republic  (  =  Madagascar):  (An- 
tsiranana),  Diego-Suarez  (=Antsiranana).  1910. 


Collector.  Dr.  Mazieres. 

Condition.  Mummy  (dried  carcass),  complete. 
Type  Series.  Holotype  only. 

Comments.  Traditionally  included  in  Triaenops  fur- 
culus,  as  in  G.  M.  Allen  (1939:  82)  and  Koopman 
(1993:  175),  but  retained  as  distinct  by  Peterson  et 
al.  (1995:  81)  pending  further  material  from  the 
area  of  the  type  locality.  Known  only  from  the  ho- 
lotype. 

Family  MORMOOPIDAE  de  Saussure, 
1860 

Genus  PTERONOTUS  Gray,  1838 

Ctiilonycteris  parnellii  pusillus  G.  M.  Allen, 

1917c 

Proc.  Biol.  Soc.  Washington,  30:  168,  23 

October. 

=  Pteronotus  parnellii  pusillus  (G.  M.  Allen, 

1917).  See  Smith  (1972:  67). 

Holotype.  MCZ  16468.  Skin  and  skull.  Female. 
Locality.    Dominican    Republic:    Santo   Domingo, 
Arroyo  Salado.  7  March  1916. 
Collector  J.  L.  Peters.  Original  number  227. 
Condition.  Skin  complete.  Skull  partial  (left  wall  of 
braincase  broken,  left  tympanic  bulla  missing). 
Type    Series.    2   paratypes;    MCZ    16599,    female; 
MCZ  16600,  female;  both  in  alcohol. 
Comments.   P.  p.  pusillus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  92)  and  Koopman  (1994: 
71). 

Ctiilonycteris  torrelG.  M.  Allen,  1916a 

Proc.  New  England  Zool.  Club,  6:  4,  8 

Februai-v. 

=  Pteronotus  quadridens  quadridens 

(Gundlach,  1840).  See  Silva-Taboada 

(1976:  7). 

Holotype.  MCZ  11672.  Body  in  alcohol,  skull  extract- 
ed. Adult  female. 

Locality.  Cuba:  (Guantanama),  Baracoa,  La  Cueva 
de  la  Majana.  15  June  1915. 

Collector  V.  J.  R.  Verrier.  Presented  to  the  MCZ 
by  Carlos  de  la  Torre. 
Condition.  Alcoholic,  skull  complete. 
Type  Series.  2  paratypes;  MCZ  11670,  male;  MCZ 
11671,  female;  both  in  alcohol. 

Family  PHYLLOSTOMIDAE  Gray,  1825 

Genus  AMETRIDA  Gray,  1847 

Ametrida  minor  H.  Allen,  1894 

Proc.  Boston  Soc.  Nat.  Hist.,  26:  240,  16 

May. 

= Ametrida  centurio  Gray,  1847.  See 

Peterson  (1965:  5). 

Holotype.  MCZ  11274.  Body  in  alcohol,  skull  extract- 
ed. Adult  male. 


144 


Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Locality.   Suriname:  Paramaribo.  Collected  some- 
time between  1832  and  1839. 
Collector.  F.  W.  Cragin. 
Condition.  Alcoholic,  skull  complete. 
Type  Series.  Holotype  only. 

Comments.  For  comments  on  the  type  locality  and 
date  of  acquisition,  see  G.  M.  Allen  (1902a:  88).  A. 
minor  actually  represents  the  male  specimens  of  A. 
centurio. 

Genus  ARTIBEUS  Leach,  1821 

Artibeus  femurvillosum  Bangs,  1899k 

Proc.  New  England  Zool.  Club,  1:  73,  24 

November. 

= Artibeus  lituratus  palmarum  J.  A.  Allen 

and  Chapman,  1897.  See  Hershkovitz 

(1949:  445). 

Holotype.  B8314.  Skin  and  skull.  Adult  male. 

Locality.  Colombia:  (La  Guajira),  La  Concepcion, 

3,000  ft  (915  m).  21  March  1899. 

Collector  W.  W.  Brown,  Jr.  Original  number  31. 

Condition.  Skin  and  skull  complete. 

Ti/pe  Series.  Holotype  only. 

Genus  EROPHYLLA  Miller,  1906 

Erophylla  sezekomi  syops  G.  M.  Allen, 

1917c 

Proc.  Biol.  Soc.  Washington,  30:  167,  23 

October. 

Holotype.  MCTj  13713.  Body  in  alcohol,  skull  extract- 
ed. Adult  male. 

Locality.  Jamaica:  (St.  James),  Montego  Bay.  14 
March  1912. 

Collector  J.  A.  Cushman. 
Condition.  Alcoholic,  skull  complete. 
Type    Series.    7   paratypes;    MCZ    13709-13712, 
13714-13716,  all  alcoholic.  MCZ  13709  and  13712 
were  sent  in  e.xchange  to  the  USNM. 
Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  171)  and  Koopman  (1994:  79). 

Genus  GLOSSOPHAGA  E.  Geoffrey, 
1818 

Glossophaga  longirostris  Miller,  1898 
Proc.  Acad.  Nat.  Sci.  Philadelj^hia,  1898, 
p.  330,  2  August. 

=  Glossophaga  longirostris  longirostris 
Miller,  1898.  See  Miller  (1913:  422). 

Holotype.  B8046.  Skin  and  skull.  Adult  female. 

Locality.    Colombia:    (Magdalena),   Santa  Marta 
Mountains,  near  Santa  Marta.  10  February  1898. 
Collector  W.  W.  Brown,  Jr.  Original  number  60. 


Condition.  Skin  complete.  Skull  partial  (left  zygo- 
matic arch  missing). 
Type  Series.  Holotype  only. 

Comments.  G.  longirostris  was  considered  a  valid 
species  by  Koopman  (1993:  184)  and  Nowak  (1999: 
368). 

Genus  LONCHOPHYLLA  Thomas,  1903 

Lonchophylla  hesperia  G.  M.  Allen,  1908 
Bull.  Mus.  Comp.  Zool,  52:  35,  July. 

Holotype.  MCZ  7011.  Body  in  alcohol,  skull  extract- 
ed. Adult  male. 

Locality.  Peru:  (Contralmirante  Villar),  Tumbes, 
Zorritos. 

Collector.  F.  H.  Bradley. 
Condition.  Alcoholic,  skull  complete. 
Type  Series.   2  paratypes;  in  the  YPM;  1034  and 
1035;  both  in  alcohol. 

Comments.  Considered  a  valid  species  by  Koop- 
man (1993:  181)  and  Nowak  (1999:  372).  This  rare- 
ly collected  bat  is  known  by  only  two  museum 
specimens  in  addition  to  the  type  series:  USNM 
283177  and  LSUMZ  14121  (Gardner  1976:  5). 

Genus  PLATYRRHINUS  6e  Saussure, 
1860 

Vampyrops  umbratus  Lyon,  1902 

Proc.  Biol.  Soc.  Washington,  15:  151,  20 

June. 

=  Platyrrhinus  umbratus  (Lyon,  1902).  See 

Koopman  (1993:  191). 

Holotype.  B8180.  Sldn  and  skull.  Adult  male. 

Locality.  Colombia:  (LaGuajira),  San  Miguel.  8 
June  1898. 

Collector  W.  W.  Brown,  Jr.  Original  number  234. 
Condition.  Sldn  and  skull  complete. 
Type  Series.  2  paratypes;  B8300,  skin,  male;  B8301, 
sldn,  male. 

Comments.  P.  umbratus  was  considered  a  valid  spe- 
cies by  Koopman  (1993:  191)  and  Nowak  (1999: 
389).  Platyrrhinus  has  priority  over  the  genus  name 
Vampyrops  (Gardner  and  Ferrell  1990:  501-503). 

Vampyrops  zarhinus  H.  Allen,  1891 
Proc.  Acad.  Nat.  Sci.  Philadelphia,  1891, 
p.  400,  22  September. 
=  Platyrrhinus  helleri  {Peters,  1866).  See 
Hall  and  Kelson  (1959:  131). 

Holotype.  MCZ  3211.  Body  in  alcohol,  skull  extract- 
ed. Adult  female,  pregnant. 

Locality.  Panama:  Canal  Zone,  Obispo.  1872.  See 
comments. 

Collector  Hassler  Expedition. 
Condition.  Alcoholic,  skull  complete. 
Type  Series.  Holotype  only. 


Type  Specimens  of  Recent  Mammals  •  Helgen  and  McFadden         145 


Comments.  In  the  original  description,  H.  Allen  re- 
port:ed  that  this  specimen  had  been  collected  in 
Brazil  by  the  Thayer  expedition.  G.  M.  Allen 
emended  this  apparently  erroneous  locality  to 
Obispo,  Panama,  in  accordance  ^A-ith  the  accession 
catalogue  of  the  MCZ  (1931:  236-237).  In  support 
of  Aliens  decision,  Rouk  and  Carter  (1972:  4)  stat- 
ed, after  examining  the  holotype  of  zarhinus,  that 
it  is  "quite  like  specimens  of  [Platyrrhiniis]  helleri 
from  Mexico  and  Central  America,  and  unlikely  to 
have  come  from  Brazil."  For  the  use  of  the  genus 
Platyrrhinus  over  Vampyrops,  see  Gardner  and 
Ferrell  (1990:  501-503). 


Genus  VAMPYRODES  Thomas,  1 900 

Vampyrodes  major  G.  M.  Allen,  1908 
Bull.  Mus.  Comp.  ZooL,  52:  38,  July. 
=  Vampyrodes  caraccioli  major  G.  M. 
Allen,  1908.  See  Handley  (1966:  766). 


Holotype.  MCZ  6756.  Body  in  alcohol.  Adult  female. 
Locality.  Panama:  San  Pablo  (now  covered  by  Ga- 
tun  Lake).  Date  unrecorded. 
Collector.  A.  Lesley. 
Condition.  Alcoholic. 
Type  Series.  Holotype  only. 

Comments.  V.  c.  major  was  retained  as  a  valid  sub- 
species by  Koopman  (1994:  88). 


Family  MOLOSSIDAE  Gervais,  1855 

Genus  MOPS  Lesson,  1842 

Chaerephon  leucostigma  G.  M.  Allen, 

1918a 

Bull.  Mus.  Comp.  ZooL,  61:  513, 

Februaiy. 

=  Mops  condylurus  leucostigma  (G.  M. 

Allen,  1918).  See  Koopman  (1994:  141). 


Holotype.  MCZ  16344.  Skin  and  skull.  Aduh  female. 
Locality.  Malagasy  Republic  (Madagascar):  (Anta- 
nanarivo), Tananarive  (=  Antananarivo).  December 
1915. 

Collector.  F.  R.  Wulsin. 

Condition.  Skin  partial  (bare  spot  on  ventrum). 
Skull  damaged  (right  and  left  zygomatic  arch  miss- 
ing; supraoccipital  chipped). 

Type  Series.  1  paratype;  MCZ  16345,  skin  and 
skull,  male. 

Comments.  Peterson  et  al.  (1995:  168)  used  the 
name  Tadarida  leucostigma. 


Mops  angolensis  orientis  G.  M.  Allen  and 

Loveridge,  1942 

Bull.  Mus.  Comp.  ZooL,  89:  166, 

Februaiy. 

=  Mops  condylurus  orientis  G.  M.  Allen 

and  Loveridge,  1942.  See  Koopman 

(1994:  141). 

Holotype.  MCZ  38829.  Skin  and  skull.  Adult  male. 
Locality.  Tanganyika  Territoiy  (=  Tanzania):  Mtwa- 
ra,  Ruviuna  River,  Kitaya,  300  ft  (92  m).  3  April 
1939. 

Collector  A.  Loveridge. 
Condition.  Skin  and  skull  complete. 
Tijpe   Series.    9   paratypes;    MCZ   38826-38828, 
38830-38835,  all  represented  by  skin  and  skull,  4 
females  and  5  males. 

Genus  OTOMOPS  Thomas,  1913 

Otomops  papuensis  Lawrence,  1948 
J.  Mammal.,  29:  413,  31  December. 

Holotype.  MCZ  45769.  Body  in  alcohol,  skull  extract- 
ed. Adult  female. 

Locality.  Papua  New  Guinea:  Vailala  River. 
Collector.  Bought  from  Ward's  Natural  Science  Es- 
tabhshment,  April  1948. 

Condition.  Alcoholic,  skull  partial  (right  zygomatic 
arch  missing). 
Type  Series.  Holotype  only. 

Comments.  Considered  a  valid  species  by  Koop- 
man (1993:  239)  and  Nowak  (1999:  482).  Accord- 
ing to  Flanneiy  (1995a:  481),  O.  papuensis  has 
been  collected  on  only  two  occasions  and,  other 
than  the  holotype,  is  known  by  only  10  specimens; 
2  in  the  BMNH,  the  remainder  in  the  biological 
collections  of  the  University  of  Papua  New  Guinea. 

Family  VESPERTILIONIDAE  Gray,  1821 

Genus  EPTES/CL/S  Rafinesque,  1820 

Eptesicus  darlingtoni  G.  M.  Allen,  1933 
J.  Mammal.,  14:  150,  15  May. 

Holotype.  MCZ  29113.  Skin  and  skull.  Adult  female. 
Locality.  Australia:  Queensland,  Queensland  Na- 
tional Park,  MacPherson  Ranges,  3,000  ft  (915  m). 
10  March  1932. 

Collector  P.  J.  Darlington,  Jr.,  Harvard  Australiaji 
Expedition.  Original  number  30. 
Condition.  Skin  and  skull  complete. 
Type  Series.  1  paratype;  MCZ  29120  (now  Queens- 
land Museum  J  5476),  skin  and  skull,  adult  female. 
Comments.  McKean  et  al.  (1978:  533)  and  Koop- 
man (1993:  203)  included  darlingtoni  in  Eptesicus 
pumilus.  However,  Koopman  also  used  the  name 
Pipistrellus  darlingtoni  (1994:   116).   Hoye  (1995: 


146         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


537)  considered  darlingtoni  to  be  a  valid  species  of 
VespadeJus,  to  which  he  gave  full  generic  rank. 

Genus  HARPIOCEPHALUS  Gray,  1842 

Harpiocephalus  rufulus  G.  M.  Allen,  1913 
Proc.  Biol.  Soc.  Washington,  26:  214,  20 
December. 

=  Harpiocephalus  harpia  rufulus  G.  M. 
Allen,  1913.  See  Ellerman  and  Morrison- 
Scott  (1951:  187). 

Holotijpe.  MCZ  14206.  Skin  and  skull.  Adult  male. 
Localitij.  Vietnam:  Tonkin,  Lao-Kai  (  =  Lao  Cai).  3 
Januaiy  1912. 

Collector.  Kobayashi  Collection.  Original  number 
14. 

Condition.  Skin  complete.  Skull  partial  (parietals 
broken). 

Type  Serie.s.  Holotype  only. 

Comments.  H.  h.  nifuliis  was  retained  as  a  valid 
subspecies  by  Koopman  (1994:  133). 

Genus  IDIONYCTERIS  Anthony,  1923 

Corynorhinus  phyllotis  G.  M.  Allen,  1916b 
Bull.  Mns.  Comp.  Zool.,  60:  352,  April. 
=  Idionycteris  phyllotis  phyllotis  (G.  M. 
Allen,  1916).  See  Tumlison  (1993:  418). 

Holotype.  MCZ  5943.  Skin  and  skull.  Adult. 

Locality.  Mexico:  San  Luis  Potosi.  24  March  1878. 
Collector.  E.  Paliner. 
Condition.  Skin  and  skull  complete. 
Tijpe  Series.  Holotype  only. 

Comments.  I.  phyllotis  was  considered  a  valid  spe- 
cies by  Koopman  (1993:  205)  and  Nowak  (1999: 
457). 

Genus  LASIURUS  Gray,  1838 

Atalapha  brachyotisJ.  A.  Allen,  1892 

Bull.  Amer.  Mus.  Nat.  Hist.,  4:  47,  25 

March. 

=  Lasiurus  borealis  brachyotis  (J.  A.  Allen, 

1892).  See  Niethammer  (1964:  595). 

Holotype.  MCZ  11143.  Body  in  alcohol.  Male. 

Locality.   (Ecuador),  Galapagos  Islands:  Chatham 
Island.  23  June  1891. 
Collector  G.  Baur. 

Condition.  Alcoholic.  The  specimen  was  received 
without  a  skull. 
Type  Series.  Holotype  only. 

Comments.  Lasiunis  brachyotis  has  often  been  ac- 
corded specific  status,  as  in  Nowak  (1999:  451).  In- 
cluded in  L.  borealis  as  a  valid  subspecies  by  Koop- 
man (1994:  129). 


Genus  /W/OT/S  Kaup,  1829 

Myotis  abbotti  nugax  G.  M.  Allen  and 

Coolidge,  1940 

Bull.  Mus.  Comp.  Zool.,  87:  137,  31 

December. 

=  Myotis  muricola  nugax  G.  M.  Allen  and 

Coolidge,  1940.  See  Koopman  (1994: 

104). 

Holotype.  MCZ  36076.  Skin  and  skull.  Adult  male. 
Locality.    Malaysia:  north  Borneo,   Sabah,   Mount 
Kinabalu,  Bundutuan,  3,500  ft  (1,068  m).  25  July 
1937. 

Collector  J.  A.  Griswold,  Jr.,  Asiatic  Primate  Ex- 
pedition. Original  number  626. 
Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  16  paratypes,  MCZ  36072-36075, 
36077-36080,  36082-83,  36085-89,  36091;  all  rep- 
resented by  skin  and  skull,  12  females  and  4  males. 

Myotis  albicinctus  G.  M.  Allen,  1919b 
J.  Mammal.,  1:  2,  28  November. 
=  Myotis  lucifugus  carissima  Thomas, 
1904.  See  Miller  and  G.  M.  Allen  (1928: 

50). 

Holotype.  MCZ  11747.  Sldn  and  skull.  Adult  male. 
Locality.  (United  States):  California,  (Tulare  Coun- 
ty), Mount  Whitney,  11,000  ft  (3,355  m).  14  July 
1915. 

Collector  G.  M.  Allen.  Original  number  1. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  1  paratyj^^e,  probably  at  USNM. 
Comments.  The  skull  of  the  holotype,  which  had 
been  mislaid  at  the  time  that  albicinctus  was  de- 
scribed, has  subsequently  been  found  and  reunited 
with  its  skin. 

Myotis  sodalis  Miller  and  G.  M.  Allen, 

1928 

Bull.  U.S.  Nat.  Mus.,  144:  130,  25  May. 

Holotype.  MCZ  10988.  Skin  and  skull.  Adult  female. 
Locality.    (United    States):    Indiana,    (Crawford 
County),  Wyandotte  cave.  7  March  1904. 
Collector  J.  O.  Sibert. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Comments.  Considered  a  valid  species  by  Koop- 
man (1993:  215)  and  Nowak  (1999:  419). 
Type  Series.  Miller  and  Allen  based  their  descrip- 
tion on  an  examination  of  443  specimens,  described 
by  locality  in  the  original  description  (1928:  133). 
30  paratypes  are  in  the  MCZ,  the  others  are  in  tlie 
FMNH,  USNM,  AMNH.  and  BMNH. 


Type  Specimens  of  Recent  Mammals  •  Hehen  and  McFadden         147 


Genus  A/VCT/CE/L/S  Rafinesque,  1819 

Nycticeius  africanus  G.  M.  Allen,  1911b 

Bull.  Mus.  Comp.  Zool.,  54:  328, 

December. 

=  Nycticeius  sctilieffeni  albiventer  Thomas 

and  Wroughton,  1908.  See  Hayman  and 

Hill  (1971:  36). 

Holotype.  MCZ  8272.  Skin  and  skull.  Male. 

Locality.  British  East  Africa  (  =  Kenya):  Meru  Riv- 
er, effluent  of  northern  Guaso  N^'iro  (  =  Ewaso  Ngi- 
ro).  11  August  1909. 

Collector.  G.  M.  Allen.  Original  number  113. 
Condition.  Skin  and  skull  complete. 
Type  Series.  Holotype  only. 

Genus  PIPISTRELLUS  Kaup,  1829 

Eptesicus  phasma  G.  M.  Allen,  1911b 

Bull.  Mus.  Comp.  Zool,  54:  327, 

December. 

=  Pipistreilus  rendalii  phasma  (G.  M.  Allen, 

1911).  See  Koopman  (1994:  117). 

Holotype.  MCZ  8279.  Skin  and  skull.  Male. 

Locality.  British  East  Africa  (  =  Kenya):  Meru  Riv- 
er, effluent  of  northern  Guaso  Nyiro  ( =  Ewaso  Ngi- 
ro).  6  August  1909. 

Collector.  G.  M.  Allen.  Original  number  94. 
Condition.  Skin  and  skull  complete. 
Type  Series.  There  is  a  small  series  of  paratypes  in 
the  MCZ. 

Scabrifer  notiusG.  M.  Allen,  1908 
Bull.  Mus.  Comp.  Zool.,  52:  46,  July. 
=  Pipistreilus  capensis  notius  (G.  M.  Allen, 
1908).  See  Koopman  (1994:  117). 

Holotype.     MCZ    4555.    Alcoholic,    skull    extracted. 
Adult  male. 

Locality.  South  Africa:  (Western  Cape),  Cape 
Town. 

Collector  Received  from  E.  L.  Lavard,  August 
1864. 

Condition.  Alcoholic,  skull  partial  (right  and  left 
zygomatic  arches  missing;  supraoccipital  chipped). 
Type  Series.  Holotyjje  only. 

Genus  PLECOTUS  E.  Geoffrey,  1813 

Plecotus  sacrimontis  G.  M.  Allen,  1908 
Bull.  Mus.  Comp.  Zool.,  52:  50,  July 
=  Piecotus  auritus  sacrimontis  G.  M.  Allen, 
1908.  See  Ognev  (1928:  607). 

Holotype.  MCZ  6932.  Body  in  alcohol.  Adult  male. 
Locality.  Japan:  (Honshu),  Mount  Fuji.  4  Decem- 
ber 1906.' 


Collector.  A.  Ovs'ston. 

Condition.  Alcoholic. 

Type  Series.  Holotype  only. 

Comments.  P.  a.  sacrimontis  was  retained  as  a  valid 

subspecies  by  Koopman  (1994:  110). 

Genus  SCOTOPHILUS  Leach,  1821 

Scotopiiiius  altilis  G.  M.  Allen,  1914d 
Bull.  Mus.  Comp.  Zool.,  58:  350,  July 
=  Scotopiiiius  leucogaster  (Gretzschmar, 
1826).  See  Koopman  (1993:  227). 

Holotype.  MCZ  14463.  Skin  and  skull.  Adult  male. 
Locality.  Sudan:  Blue  Nile,  north  of  (Er)  Roseires, 
Aradeiba.  22  January  1913. 

Collector  G.  M.  Allen,  Phillips  Sudan  Expedition. 
Original  number  7.3. 
Condition.  Skin  and  skull  complete. 
Type   Series.   3  paratypes;   MCZ   14462,  skin  and 
skull,  male,  exchanged  to  FMNH;  and  14610  and 
14611,  both  males  in  alcohol. 

Family  THYROPTERIDAE  Miller,  1907 

Genus  THYROPTERA  Sp\x,  1823 

Thyroptera  tricolor  albigula  G.  M.  Allen, 

1923c 

Proc.  New  England  Zool.  Club,  9:  1,  10 

December. 

=  Thyroptera  tricolor  albiventer  (Tomes, 

1856).  See  Dunn  (1931:  430). 

Holotype.  MCZ  20143.  Body  in  alcohol,  skull  extract- 
ed. Adult  female. 

Locality.  Panama:  Gutierrez,  25  miles  (40.2  km) 
inland  from  Chiriquiscito  on  trail  from  Chiriqui  La- 
goon, Bocas  del  Toro  to  Boquete,  Chiriqui.  August 
1923. 

Collector  E.  R.  Dunn  and  C.  B.  Dvuyea. 
Condition.  Alcoholic,  skull  complete. 
Type  Series.  3  paratypes;  MCZ  20144,  adult  male; 
MCZ  20145,  juvenile;  MCZ  20146,  juvenile;  all  in 
alcohol. 

Order  ARTIODACTYLA  Owen,  1848 

Family  TAYASSUIDAE  Palmer,  1897 

Genus  PEC/\R/ Reichenbach,  1835 

Tayassu  crusnigrum  Bangs,  1 902b 
Bull.  Mus.  Comp.  Zool.,  39:  20,  April. 
^Pecari  tajacu  crusnigrum  (Bangs,  1902). 
See  Hershkovitz  (1951:  567). 

Holotype.  MCZ  10163.  Skin  and  skull.  Adult  male. 
Locality.   Panama:  Chiriqui,  Boquete  (  =  Ba)o  Bo- 
quete), 4,000  ft  (1,220  m).  13  April  1901. 


148         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Collector.  W.  W.  Brown,  Jr.  Original  number  290. 
Condition.  Skin  and  skull  complete. 
Type   Series.   2  paratypes;   MCZ   10162,  adult  fe- 
male; MCZ  10164,  juvenile  female. 
Comments.  P.  t.  crusnignim  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  1080)  under  the  genus 
Dicotijles. 

Tayassu  torvus  Bangs,  1 898k 

Proc.  Biol.  Soc.  Washington,  12:  164,  10 

August. 

=  Pecari  tajacu  torvus  (Bangs,  1898).  See 

comments. 

Holotype.  B8038.  Sldn  and  skull.  Adult  male. 

Locality.  Colombia:  Magdalena,  Santa  Marta.  26 
January  1898. 

Collector.  W.  W.  Brown,  Jr.  Original  number  50. 
Condition.  Skin  and  skull  complete. 
Type  Series.  Holotype  only. 

Comments.  Retained  as  a  valid  subspecies  by  Ca- 
brera (1961:  319)  under  the  genus  Tayassu.  Use  of 
the  genus  Pecari  follows  Grubb  (1993:  380). 

Family  MONODONTIDAE  Gray,  1821 

Genus  DELPHINAPTERUS  Lacepede, 
1804 

Beluga  declivis  Cope,  1865 

Proc.  Acad.  Nat.  Sci.  Philadelphia,  17: 

278. 

=  Delphinapterus  leucas  (Pallas,  1776). 

See  Hershkovitz  (1966:  111). 

Holotype.  MCZ  1195.  Skull  and  postcranial  skeleton. 
Locality.  "Arctic  Seas"  (probably  Greenland). 
Collector.  E.  K.  Kane. 

Condition.   Skull  partial  (right  mandibular  ramus 
missing).  Postcranial  skeleton  complete  except  for 
missing  left  flipper.  Right  flipper  and  tail  are  un- 
cleaned,  with  tissue  largely  intact. 
Type  Series.  Holotype  only. 

Comments.  Hershkovitz  (1966:  111)  erroneously 
stated  that  the  holotype  of  declivis  was  deposited 
in  the  Academy  of  Natural  Sciences  in  Philadel- 
phia, and  the  holotype  of  Beluga  concreta  Cope, 
1865  was  in  the  MCZ.  The  opposite  is  in  fact  true; 
Philadelphia  holds  the  type  of  B.  concreta. 

Family  PHOCOENIDAE  Gray,  1825 

Genus  NEOPHOCAENA  Palmer,  1899 

Neomeris  asiaeorientalis  Pilleri  and  Gihr, 

1972 

Invest.  Cetacea,  4:  126. 

=  Neophocaena  phocaenoides 

asiaorientalis  (Pilleri  and  Gihr,  1972).  See 

van  Bree  (1973:  17). 

Holotype.  MCZ  19998  (but  see  comments).  Skull  and 
postcranial  skeleton.  Adult  male. 


Locality.  China:  Kiangsu  (=Jiangsu),  Kiangyin,  80 

miles    (129   km)   northwest   of  Shanghai.    7  April 

1922. 

Collector  F.  R.  Wulsin. 

Condition.  Skin  and  postcrajiial  skeleton  complete. 

Type  Series.  Holotype  only. 

Comments.  For  a  discussion  of  the  nomenclature 

and    synonymy    of  this    form,    consult   van    Bree 

(1973).  Because  Neomeris  asiaorientalis  Pilleri  and 

Gihr,  1972  is  in  fact  a  replacement  name  for  the 

preoccupied  name  Delpliinus  inelas  Schlegel,  1841, 

the  holotype  of  this  new  name  is  the  same  as  that 

of  Schlegel's  name,  RMNH  23079. 

Family  CERVIDAE  Goldfuss,  1820 

Genus  ODOCO//.EL/S  Rafinesque,  1832 

Cariacus  osceola  Bangs,  1896b 

Proc.  Biol.  Soc.  Washington,  10:  26,  25 

February. 

=  Odocoileus  virginianus  osceola  (Bangs, 

1896).  See  Lydekker  (1915:  148). 

Holotype.  B2394.  Skin  and  skull.  Adult  female. 
Locality.   (United  States):  Florida,  Citrus  County, 
Citronelle.  29  December  1893. 
Collector.  F.  L.  Small.  Original  number  1107. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Ttjpe  Series.  4  paratypes;  B2391,  adult  male, 
B2392,  adult  male,  B2393,  adult  female,  B2395, 
juvenile  male,  all  represented  by  skin  and  skull. 
Comments.  O.  v.  osceola  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  1096)  and  Smith  (1991: 
1). 

Odocoeleus  [sic]  virginianus  louisianae  G. 

M.  Allen,  1901 

Amer.  Nat.,  35:  449.  28  June. 

=  Odocoileus  virginianus  macroura 

(Rafinesque,  1817).  See  Miller  and 

Kellogg  (1955:  804). 

Holotype.  B9111.  Skin  and  skull.  Adult  male. 

Locality.    (United   States):    Louisiana,    Morehouse 
Parish,  Mer  Rouge.  8  November  1898. 
Collector  B.  V.  Lilly 

Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  3  paratypes;  B9112,  B8622,  B8623,  all 
males  represented  by  skin  and  skull. 

Odocoileus  americanus  borealis  Miller, 

1900 

Bull.  New  York  State  Mus.  8:  83,  21 

November. 

=  Odocoilus  virginianus  borealis  Miller, 

1900.  See  Trouessart  (1905:  704). 

Holotype.  B4999.  Skin  and  skufl.  Adult  male. 

Locality.  (United  States):  Maine,  (Hancock  Coun- 
ty), Bucksport.  12  December  1895. 


Type  Specimens  of  Recent  Mammals  •  Helaen  and  McFadden 


149 


Collector.  A.  G.  Dorr. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Holotype  only. 

Comments.  O.  v.  horealis  was  retained  as  a  valid 
subspecies  bv  Hall  (1981:  1092)  and  Smith  (1991: 
1). 

Odocoileus  virginianus  clavium  Barbour 

and  G.  M.  Allen,  1922 

J.  Mammal.,  3:  73,  9  May. 

Holotype.    MCZ  19120.   Skull  and  head  skin.  Adult 
male. 

Locality.  (United  States):  Florida,  (Monroe  Coun- 
ty), Big  Pine  Key  Winter  1920. 
Collector  T.  Barbour. 

Condition.  Skin  and  skaill  complete.  Mandible  dis- 
articulated. 

Type  Series.  2  paratyjDcs,  MCZ  18497,  sldn  and 
skull,  juvenile  male;  MCZ  18060,  sldn  and  skull, 
juvenile  male. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  1093)  and  Smith  (1991:  1). 

Genus  R/\A/G/FE/=?  Hamilton  Smith,  1827 

Rangifer  arcticus  caboti  G.  M.  Allen, 

1914a 

Proc.  New  England  Zool.  Club,  4:  104, 

24  March. 

=  Rangifer  tarandus  caribou  (Gmelin, 

1788).  See  Banfield  (1962:  70). 

Holotype.  MCZ  15372.  One  shed  antler.  Adult  male. 
Locality.    Canada:    northeast    coast    of   Labrador, 
about  30  miles  (48.3  km)  north  of  Nachvak.  1909. 
Collector  O.  Biyant. 
Condition.  Single  antler;  complete. 
Type  Series.  Holotype  only. 

Rangifer  terraenovae  Bangs,  1896! 

Preliminaiy  Description  of  the 

Newfoundland  Caribou,  Boston,  p.  1,  11 

November. 

=  Rangifer  tarandus  caribou  (Gmelin, 

1788).  See  Banfield  (1962:  70). 

Holotype.  B3778.  Skull  and  head  sldn.  Adult  male. 
Locality.  Canada:  Newfoundland,  Codroy  8  Sep- 
tember 1895 
Collector  E.  Doane. 

Condition.  Skull  and  head  sldn  complete. 
Type  Series.  Bangs  remarks  that  he  has  "secured  a 
series  of  this  fine  caribou,"  corresponding  to 
B3779-B3781  and  B5757-B5760. 
Comments.  J.  A.  Allen  published  a  description  of 
Rangifer  terraenovae  on  21  November  1896,  spec- 
ifying AMNH    11775,   a  mounted  specimen  of  a 


male  adult,  as  the  holotype  (1896:  233).  Bangs'  de- 
scription of  this  taxon  pre-dates  J.  A.  Allen's  by  10 
days  and  thus  has  priority. 

Family  BOVIDAE  Gray,  1821 

Genus  DAIVIALISCUS  Sclater  and 
Thomas,  1894 

Damaliscus  phillipsi  Harper,  1 939 

Proc.  Biol.  Soc.  Washington,  50:  90,  5 

June. 

=  Damaliscus  pygargus  phiillipsi  Harper, 

1939.  See  comments. 

Holotype.  MCZ  35443.  Skull  and  skin.  Adult  male. 
Locality.  South  Africa:  Orange  Free  State.  23  July 
1935. 

Collector.  P.  Andreka.  Original  number  1958d. 
Condition.    Skin  complete.   Skull  partial  (most  of 
palate,    left    maxilla,    and    left    mandibular   ramus 
missing). 

Type  Series.  Paratype  material  consists  of  MCZ 
35444,  skin  and  skull  of  an  adult  female  as  well  as 
the  following,  which  Haij)er  examined  in  the  col- 
lection of  the  Academy  of  Natural  Sciences  of  Phil- 
adelphia; "a  mounted  head,  a  skull,  and  a  set  of 
horns  purchased  in  Kimberley,  Cape  Province;  a 
mounted  head  and  a  set  of  horns  from  'South  Af- 
rica'; and  two  skins  and  skulls  from  the  Zoological 
Society  of  Philadelphia." 

Comments.  Ansell  (1972;  55)  used  the  name  Dam- 
aliscus dorcas  phillipsi.  For  the  use  of  pygargns 
over  dorcas,  see  Rookmaaker  (1991;  190). 

Order  CARNIVORA  Bowdich,  1821 

Family  CANIDAE  Fischer  de  Waldheim, 
1817 

Genus  CANIS  Linnaeus,  1758 

Canis  lupus  beothiucus  G.  M.  Allen  and 

Barbour,  1937 

J.  Mammal,  18:  230,  14  May. 

Holotype.   MCZ  351.  Skull  and  postcranial  skeleton. 
Adult,  probably  male. 

Locality.  Canada:  Newfoundland.  About  1865. 
Collector  J.  M.  Nelson. 

Condition.  Skull  and  postcranial  skeleton  com- 
plete. 

Type  Series.  4  paratypes;  348,  skull,  adult  male; 
349,  skull,  adult,  probably  male;  350,  skull,  adult 
female;  MCZ  28726,  skin. 

Comments.  C.  I.  beothiicus  became  extinct  around 
1911.  Retained  as  a  vahd  subspecies  by  Hall  (1981: 
930). 


150         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Pachycyon  robustus  J.  A.  Allen,  1885 

Mem.  Mus.  Comp.  Zool.,  10:  4, 

December. 

=  Canis  familiaris  Linnaeus,  1758.  See  G. 

M.  Allen  (1920a:  498). 

Holotype.  MCZ  7091.  Postcranial  skeleton. 

Locality.  (United  States):  Virginia,  Lee  County,  Ely 
Cave.  Probably  1875. 
Collector.  N.  S.  Shaler. 

Condition.  Skeleton  partial  (right  scapula,  right  hu- 
merus, right  femur,  right  tibia,  pelvis). 
Type  Series.  Holotype  only. 

Comments.  This  specimen  is  a  domesticated  dog  of 
Native  Americans.  C.  lupus  familiaris  is  the  name 
now  widely  used  for  the  domestic  dog  (Wozencraft 
1993:  281).  P.  robustus  is  the  type  species  of  Pa- 
chycyon ].  A.  Allen,  1885.  Pachycyon  is  a  synonym 
oiCanis  Linnaeus,  1758,  which  is  commonly  over- 
looked, for  example,  in  Wozencraft  (199.3)  and  Mc- 
Kenna  and  Bell  (1997). 

Genus  CERDOCYON  Hamilton  Smith, 
1839 

Cerdocyon  thous  germanus  G.  M.  Allen, 

1923b 

Proc.  Biol.  Soc.  Washington,  36:  55,  28 

March. 

Holotype.  MCZ  19850.  Skin  and  skull.  Adult. 

Locality.  Colombia:  high  savannali  of  Bogota,  9,000 
ft  (2,745  m). 

Collector  N.  Maria.  Original  number  25. 
Condition.  Skin  and  skull  complete. 
Type   Series.   5  paratypes;   MCZ   19849,  skin  and 
skull,  juvenile;  MCZ  20097,  sldn  and  skull,  juvenile 
male;  3  specimens  from  the  AMNH  are  also  men- 
tioned in  the  description. 

Comments.  Retained  as  a  valid  subspecies  by  Berta 
(1982:  1). 

Urocyon  aquilus  Bangs,  1 898h 

Proc.  Biol.  Soc.  Washington,  12:  93,  30 

April. 

=  Cerdocyon  thous  aquilus  (Bangs,  1898). 

See  Langguth  (1969:  178). 

Holotype.  B8001.  Skin  and  skull.  Adult  male. 

Locality.  Colombia:  (Magdalena),  Santa  Marta 
Mountains,  between  2,000  and  3,000  ft  (610-915 
m).  10  February  1898. 

Collector.  W.  W.  Brown,  Jr.  Original  number  58. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  1  paratype;  B8002,  skin  and  skull, 
adult  female. 

Comments.  Retained  as  a  valid  subspecies  by  Berta 
(1982:  1). 


Genus  L/ROC/OA/ Baird,  1858 

Urocyon  cinereoargenteus  furvus  G.  M. 

Allen  and  Barbour,  1923 

Bull.  Mus.  Comp.  Zool,  65:  266, 

February. 

Holotype.  MCZ  19774.  Skin  and  skull.  Probably  fe- 
male. 

Locality.   Panama;  Canal  Zone,  3  miles  (4.8  km) 
west  of  Balboa.  April  1922. 
Collector  T.  Barbour  and  W.  S.  Brooks. 
Condition.  Sldn  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Holotype  only. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  943)  and  Fritzell  and  Haroldson  (1982;  1). 

Urocyon  cinereoargenteus  ocyttious 

Bangs,  1899h 

Proc.  New  England  Zool.  Club,  1:  43,  5 

June. 

Holotype.  B4290.  Skin  and  skull.  Adult  female. 

Locality.  (United  States):  Wisconsin,  Grant  Coun- 
ty, Platteville.  25  Januaiy  1896. 
Collector  N.  E.  France. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Holotype  only. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  943)  and  Fritzell  and  Haroldson  (1982:  1). 

Genus  UL/LPES  Frisch,  1775 

Vulpes  deletrix  Bangs,  1 898d 

Proc.  Biol.  Soc.  Washington,  12:  36,  24 

March. 

=  Vulpes  vulpes  rubricosa  Bangs,  1 898. 

See  Churcher  (1960:  359). 

Holotype.  B6967.  Sldn  and  skull.  Adult  female. 
Locality.     (Canada):     Newfoundland,     Bay     St. 
George.  24  April  1897. 
Collector  E.  Doane. 
Condition.  Sldn  and  skull  complete. 
Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Vulpes  pennsylvanica  vafra  Bangs,  1 897f 
Proc.  Biol.  Soc.  Washington,  11:  53,  16 
March. 

Name  preoccupied  by  Vulpes  vafer 
Leidy,  1869. 


Type  Specimens  of  Recent  Mammals  •  Helpen  and  McFadden 


151 


Vulpes  pennsylvanica  rubricosa  Bangs, 

1898a 

Science,  n.  sen,  7:  271,  25  Februaiy. 

(Replacement  name  for  Vulpes 

pennsi/Ivanica  vafra  Bangs,  1897) 

=  Vulpes  vulpes  rubricosa  Bangs,  1898. 

See  Churcher  (1960:  359). 

Holotype.  B116.  Skin  and  skull.  Adult  female. 

Locality.  (Canada):  Nova  Scotia,  Digby.  3  Novem- 
ber 1893. 

Collector.  O.  Bangs. 
Condition.  Skin  and  skull  complete. 
Type  Series.  4  paratypes;  B1991,  skin  and  skull,  and 
B2001,  skull,  both  adult  males;  B1992,  skin  and 
sk-ull,  and  B2002,  skull. 

Comments.  V.  v.  rubricosa  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  939).  Vulpes  fulvus  nib- 
ricatu.s  Miller,  1900  (p.  128)  was  a  misspelling  and 
thus  accidental  renaming  of  Vulpes  pennsylvanica 
rubricosa  Bangs,  1898. 

Vulpes  rubricosa  bangsi  Merriam,  1900 
Proc.  Washington  Acad.  Sci.,  2:  667,  28 
December  28. 

=  Vulpes  vulpes  rubricosa  Bangs,  1898. 
See  Churcher  (1960:  359). 

Holotype.  B8880.  Skin  and  skull.  Juvenile  female. 
Locality.  (Canada):  Labrador,  Lance  (  =  Lanse)  au 
Loup.  2  October  1899. 
Collector  E.  Doane. 

Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  1  paratype;  B8879,  skin  and  skull, 
adult  male. 

Family  URSIDAE  Fischer  de  Waldheim, 
1817 

Genus  L/RSUS  Linnaeus,  1758 

Ursus  (Euarctos)  americanus  sornborgeri 

Bangs,  1898j 

Amer.  Nat.,  32:  500,  July. 

=  Ursus  americanus  americanus  Pallas, 

1780.  See  Bangs  (1909:  467). 

Holotype.  B7411.  Skull.  Adult,  probably  female. 
Locality.  Canada:  Labrador,  Okkak  (  =  Okak).  Sum- 
mer 1897. 

Collector  J.  D.  Sornborger,  obtained  "from  the  Es- 
kimo." 

Condition.  Skull  complete. 

Type  Series.  2  paratypes;  B7412,  skull,  female; 
B7413,  skull,  female. 

Comments.  A  skull  from  Hopedale,  Labrador 
(MCZ  7365),  has  in  the  past  been  erroneously  la- 
beled as  the  holotype  of  sornborgeri;  B7411,  the 


true  holoty[3e  of  sornborgeri,  is  now  correctly  la- 
beled as  such. 

Family  PROCYONIDAE  Gray,  1825 

Genus  PROCYON  Storr,  1780 

Procyon  gloveralleni  Nelson  and  Goldman, 

1930 

J.  Mammal.,  11:  453,  11  November. 

=  Procyon  /otor  (Linnaeus,  1758).  See 

Corbet  and  Hill  (1991:  104). 

Holotype.  MCZ  18591.  Skin  and  skull.  Juvenile  male. 
Locality.  Barbados.  1920. 
Collector  F.  Watts. 
Condition.  Skin  and  skull  complete 
Type  Series.  Holotype  only. 

Comments.  Considered  a  valid  species  by  Wozen- 
craft  (1993:  335)  and  Nowak  (1999:  698)  but  almost 
certainly  introduced  to  Barbados  in  the  17th  cen- 
tuiy  (Helgen  and  Wilson,  in  prep.).  The  last  rac- 
coon on  Barbados  was  seen  in  1964,  and  the  pop- 
ulation is  probably  extinct. 

Procyon  lotor  elucus  Bangs,  1898b 
Proc.  Boston  Soc.  Nat.  Hist.,  28:  219,  15 
March. 

Holotype.  B3502.  Skin  and  skull.  Adult  male. 

Localitij.  (United  States):  Florida,  Brevard  County, 
Oak  Lodge,  east  peninsula  opposite  Micco.  15  Feb- 
ruary 1895. 
Collector.  O.  Bangs. 

Condition.   Skin  complete.  Skull  partial  (condyle, 
coronoid,  and  angular  processes  of  left  mandibular 
ramus  broken).  Mandible  disarticulated. 
Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 
(1981:  968). 

Procyon  maynardi  Bangs,  1898g 

Proc.  Biol.  Soc.  Washington,  12:  92,  30 

April. 

=  Procyon /otor(Linneaus,  1758).  See 

Koopman  et  al.  (1957:  164). 

Holotype.  B7750.  Skin  and  skull.  Juvenile  male. 
Locality.   Bahamas:  New  Providence  Island,  Nas- 
sau. August  1897. 
Collector  H.  L.  Claridge. 

Condition.  Skin  complete.  Skull  partial  (broken 
from  frontals  to  occiput).  Mandible  disarticulated. 
Type  Series.  Holotype  only. 

Comments.  Considered  a  valid  species  by  Wozen- 
craft  (1993:  336)  and  Nowak  (1999:  698)  but  un- 
doubtedly a  recent  introduction  to  New  Providence 
Island  (see  Olson  and  Pregill,  1982:  5). 


152         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Family  MUSTELIDAE  Fischer  de 
Waldheim,  1817 

Genus  LONTRA  Gray,  1843 

Lutra  degener  Bangs,  1 898d 

Proc.  Biol.  Soc.  Washington,  12:  35,  24 

March. 

=  Lontra  canadensis  canadensis 

(Sclireber,  1776).  See  van  Zyll  de  Long 

(1972:  81). 

Holotype.  B6965.  Sldn  and  skull.  Adult  male. 

Locality.  Canada:  Newfoundland,  Bay  St.  George. 

23  April  1897. 

Collector.  E.  Doane. 

Condition.  Skin  and  skull  complete. 

Type  Series.  Paratype  material  consists  of  B6966, 

skin  and  skull  of  an  adult  female,  mentioned  by 

number  in  the  original  description,  as  well  as  "two 

extra  skulls,"  corresponding  to  B3755  and  B3799, 

and  "a  large  series  of  unsexed  otter  skulls  from 

Newfoundland,"  corresponding  to  MCZ  494—508. 

Lutra  hudsonica  vaga  Bangs,  1 898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  224,  15 

March. 

=  Lontra  canadensis  laxatina  F.  Cuvier, 

1823.  See  van  Zyll  de  Long  (1972:  81). 

Holotype.  B5749.  Skin  and  skull.  Adult  male. 

Locality.  (United  States),  Florida,  Brevard  Count)-, 

Micco.  17  March  1897. 

Collector  F.  R.  Hunter. 

Condition.  Skin  and  skull  complete. 

Type  Series.   4  paratypes;   B4995,  skin  and  skull, 

adult  female;  B4998,  sldn  and  skull,  adult  male; 

B6092,  skin  and  skull,  adult  male;  B6093,  sldn  and 

skull,  adult  female. 

Genus  MARTES  Pinel,  1792 

Mustela  atrata  Bangs,  1897b 

Amer.  Nat.,  31:  162,  1  February. 

=  Martes  americana  atrata  (Bangs,  1897). 

See  G.  M.  Allen  (1942:  166) 

Holotype.  B5752.  Skin  and  skull.  Adult  female. 

Locality.  Canada:  Newfoundland,  Bay  St.  George. 
29  September  1896. 

Collector  E.  Doane.  Original  number  2. 
Condition.  Sldn  and  skull  complete. 
Type  Series.   12  paratypes;  B5751,  skin  and  skull, 
adult    female;    MCZ    492-93,    509-517,    unsexed 
skulls. 

Comments.  M.  a.  atrata  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  983). 


Mustela  brumalis  Bangs,  1898j 

Amer.  Nat.,  32:  502,  July 

=  Martes  americana  atrata  (Bangs,  1897). 

See  Clark  et  al.  (1987:  1). 

Holotype.  B7417.  Skull.  Adult,  probably  male. 

Locality.  Canada:  Labrador,  Okkak  (  =  Okak).  Sum- 
mer 1897. 

Collector  J.  D.  Sornborger,  obtained  "from  the  Es- 
kimo." 

Condition.  Skull  complete. 

Type    Series.    2    paratypes;    B7418,    skull;    B7419, 
skull;  both  probably  male. 

Genus  MUSTELA  Linnaeus,  1758 

Mustela  cicognanii  mortigena  Bangs,  1913 
Bull.  Mus.  Comp.  ZooL,  54:  511,  July 
=  Mustela  erminea  rictiardsonii  Bonaparte, 
1838.  See  Hall  (1951:  110). 

Holotype.  B3745.  Skin  and  skull.  Adult  male. 

Locality.  Canada:  Newfoundland,  Bay  St.  George. 

27  September  1895. 

Collector.  E.  Doane.  Original  number  1. 

Condition.  Skin  and  skull  complete. 

Type  Series.  There  is  a  series  of  paratypes  in  the 

MCZ. 

Putorius  frenatus  neomexicanus  Barber 

and  Cockerell,  1898 

Proc.  Acad.  Nat.  Sci.  Philadelphia,  1898, 

p.  188,  May 

=  Mustela  frenata  neomexicana  (Barber 

and  Gocl<erell,  1898).  See  Miller  (1912: 

100). 

Holotype.  MCZ  10475.  Skin  with  extra  tail,  and  skull. 
Adult  male. 

Locality.  (United  States):  New  Mexico,  (Dona  Ana 
County),  Mesilla,  shore  of  Armstrongs'  Lake,  3,800 
ft  (1,159  m).  1  Februaiy  1898. 
Collector  A.  C.  Tyson.  Original  number  58. 
Condition.  Sldn  complete,  but  tail  poorly  prepared. 
The  tail  belonging  to  a  discarded  topotype  is  tied 
to  the  holotype  as  an  example.  Skull  complete. 
Type  Series.  A  topotype  taken  at  the  same  time  as 
the  holotype  was  partially  decomposed  and  dis- 
carded; the  tail  of  this  specimen  is  included  with 
MCZ  10475.  The  original  description  also  refers  to 
"a  specimen,  without  any  histoiy,  in  alcohol  ...  in 
the  collection  of  the  New  Mexico  Agricultural  Col- 
lege." 

Comments.   M.  f.   neomexicana  was  retained  as  a 
valid  subspecies  by  Hall  (1981:  995). 


Type  Specimens  of  Recent  Mammals  •  Helpen  and  McFadden        153 


Putorius  (Arctogale)  longicauda  oribasus 

Bangs,  1899m 

Proc.  New  England  Zool.  Club,  1:  81,  27 

December. 

=  Mustela  frenata  oribasus  (Bangs,  1899). 

See  Hall  (1936:  105). 

Holotype.  B9058.  Skin  and  skull.  Adult  female. 

Locality.  Canada:  British  Cokmibia,  source  of  Ket- 
tle River,  7,500  ft  (2,288  m).  10  September  1898. 
Collector.  A.  C.  Brooks.  Original  number  1368. 
Conclition.  Skin  and  skull  complete. 
Type  Series.  Holotype  only. 

Comments.  M.  f.  oribasus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  998). 

Putorius  (Lutreola)  lutensis  Bangs,  1 898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  229,  15 

March. 

=  Musteia  vison  lutensis  (Bangs,  1898). 

See  Hollister  (1913:  474). 

Holotype.  B7225.  Skin  and  skull.  Adult  male. 

Localitij.  (United  States),  Florida,  St.  Johns  Coun- 
ty, salt  marsh  opposite  Matanzas  Inlet.  16  February 
1897. 

Collector.  O.  Bangs.  Original  number  7. 
Condition.  Skin  and  skull  complete. 
Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  M.  v.  lutensis  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  1003)  and  Lariviere 
(1999:  1). 

Putorius  (Arctogale)  muricus  Bangs,  1 899j 

Proc.  New  England  Zool.  Club,  1:  71,  31 

July. 

=Mustela  erminea  muricus  (Bangs,  1899). 

See  Hall  (1945:  77). 

Holotype.  B9146.  Skin  and  skull.  Juvenile  male. 
Locality.    (United   States):   California,   El   Dorado 
County,  Echo,  7,500  ft  (2,288  m).  15  July  1897. 
Collector  W.  W.  Price  and  E.  M.  Nutting.  Original 
number  266. 

Conclition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  Holotype  only. 

Comments.  M.  e.  muricus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  990)  and  King  (1983:  1). 

Putorius  noveboracensis  notius  Bangs, 

1899i 

Proc.  New  England  Zool.  Club,  1:  53,  9 

June. 

=  Mustela  frenata  noveboracensis 

(Emmons,  1840).  See  Hall  (1936:  104). 

Holotype.  B2678.  Skin  and  skull.  Juvenile  male. 
Locality.    (United   States):    North   Carolina,    Bun- 
combe County,  Weavei"ville.  10  July  1892. 


Collector.  J.  S.  Cairns.  Original  number  2214. 
Condition.  Sldn  complete.  Skull  partial  (two  frag- 
ments only,  premaxilla— lachrymal).  Mandible  dis- 
articulated. 

Type  Series.  2  paratypes;  AMNH  1247,  adult  male; 
USNM  32239,  adult  male. 

Putorius  occisor  Bangs,  18991 

Proc.  New  England  Zool.  Club,  1:  54,  9 

June. 

=  Mustela  frenata  occisor  {Bangs,  1899). 

See  Hall  (1936:  104). 

Holotype.  B9102.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Maine,  Hancock  County, 
Bucksport,  near  mouth  of  Penobscot  River.  15  Jan- 
uary 1899. 

Collector  A.  G.  Dorr. 

Condition.  Skin  complete.  Skull  slightly  damaged 
(left  zygomatic  arch  broken). 

Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  M.  f.  occisor  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  997). 

Putorius  rixosus  Bangs,  1896a 

Proc.  Biol.  Soc.  Washington,  10:  21,  25 

Februaiy. 

=  Mustela  nivalis  rixosa  (Bangs,  1896). 

See  Reichstein  (1958:  169). 

Holotype.  B642.  Skin  and  skull.  Adult  female. 

Locality.  Canada:  Saskatchewan,  Osier  15  July 
1893. 

Collector  W.  C.  Colt.  Original  number  79/181. 
Condition.   Skin  complete.  Skull  slightly  damaged 
(left  zygomatic  arch  broken). 

Type  Series.  Three  specimens  other  than  the  ho- 
lotype are  mentioned  by  number  in  the  original 
description;  MCZ  5532,  USNM  4231,  probably  fe- 
male, and  USNM  13904,  probably  male.  All  are 
unsexed  skins. 

Comments.  M.  n.  rixosa  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  993)  and  Sheffield  and  King 
(1994:  1). 

Putorius  vison  energumenos  Bangs, 

1896c 

Proc.  Boston  Soc.  Nat.  Hist.,  27:  5, 

March. 

=  Mustela  vison  energumenos  (Bangs, 

1896).  See  Miller  (1912:  101). 

Holotype.  B3555.  Skin  and  skull.  Adult  male. 

Locality.  Canada,  British  Columbia,  Sumas.  23 
September  1895. 

Collector  A.  C.  Brooks.  Original  number  514. 
Condition.  Skin  and  skull  complete. 


154         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Type  Series.  1  paratype;  B3556,  skin  and  skull,  ju- 
venile male. 

Comments.  M.  v.  energumenos  was  retained  as  a 
valid  subspecies  by  Hall  (1981:  1001)  and  Lariviere 
(1999:  1). 

Putorius  (Lutreola)  vulgivagus  Bangs, 

1895b 

Proc.  Boston  Soc.  Nat.  Hist.,  26:  539,  31 

July. 

=  Mustela  vison  vulgivaga  (Bangs,  1895). 

See  Miller  (1912:  102). 

Holotype.  B2751.  Skin  and  skull.  Adult  male. 

Locality.   (United  States):  Louisiana,  Plaquemines 
Parish,  Burbridge.  10  Januaiy  1895. 
Collector.  F.  L.  Small.  Original  number  1439/54. 
Condition.  Skin  and  skull  complete. 
Tijpe  Series.   10  paratypes;  B2752-B2761,  all  rep- 
resented by  sldn  and  skull,  9  males  and  1  female. 
Comments.  M.  v.  vulgivaga  was  retained  as  a  valid 
subspecies    by    Hall    (1981:    1004)    and    Lari\'iere 
(1999:  1). 

Putorius  xanthogenys  mundus  Bangs, 

1899i 

Proc.  New  England  Zool.  Club,  1:  56,  9 

June. 

=  Mustela  frenata  munda  (Bangs,  1899). 

See  Hall  (1936:  107). 

Holotype.  B5459.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  California,  Marin  Coun- 
ty, Point  Reyes.  19  June  1896. 
Collector  C.  A.  Allen.  Original  number  931. 
Condition.   Skin  complete.  Skull  slightly  damaged 
(left  zygomatic  arch  broken). 

Type  Series.  1  paratype,  B8632  (not  B8631,  men- 
tioned erroneously  in  the  original  description),  skin 
and  skull,  male. 

Comments.  M.  f.  munda  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  995). 

Family  MEPHITIDAE  Bonaparte,  1845 

Genus  MEPHITIS  E.  Geoffrey  and  G. 
Cuvier,  1795 

Mephistis  avia  Bangs,  1898c 

Proc.  Biol.  Soc.  Washington,  12:  32,  24 

March. 

=- Mephitis  mephitis  avia  Bangs,  1898.  See 

Hall  (1936:  65). 

Holotype.  B5747.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Illinois,  Mason  County, 
San  Jose.  10  March  1897. 

Collector.  H.  H.  and  C.  S.  Brimley  Original  num- 
ber 2500. 


Condition.  Skin  and  skull  complete. 
Ti/pe   Series.    1   paratype;   B5783,   skin   and  skull, 
adult  male. 

Comments.  M.  m.  avia  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  1019)  and  Wade-Smith  and 
Verts  (1982:  1). 

Mephitis  mephitica  elongata  Bangs,  1895b 

Proc.  Boston  Soc.  Nat.  Hist.,  26:  531,  31 

July. 

=  Mephitis  mephitis  elongata  Bangs,  1895. 

See  A.  H.  Howell  (1921:  39). 

Holotype.  B3051.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Florida,  Brevard  County, 

Micco.  5  March  1895. 

Collector  O.  Bangs. 

Condition.  Skin  and  skull  complete. 

Type  Series.  There  is  a  series  of  paratypes  in  the 

MCZ. 

Comments.  Retained  as  a  valid  subspecies  by  Hall 

(1981:  1019)  and  Wade-Smith  and  Verts  (1982:  1). 

Mephitis  mephitica  scrutator  Bangs,  1 896i 

Proc.  Biol.  Soc.  Washington,  10:  141,  28 

December. 

=  Mephitis  mephitis  mesomelas 

Lichtenstein,  1832.  See  Hall  (1936:  66). 

Holotype.  B2889.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Louisiana,  Acadia  Parish, 
Caitville.  25  May  1895. 

Collector  F.  L.  Small.  Original  number  1842. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Ti/pe  Series.  1  parats-pe,  B2886,  skin  and  skull, 
adult  female. 

Mephitis  spissigrada  Bangs,  1898c 

Proc.  Biol.  Soc.  Washington,  12:  31,  24 

March. 

=  Mephitis  mephitis  spissigrada  Bangs, 

1898.  See  Hall  (1936:  67). 

Holotype.  B3699.  Skin  and  skull.  Adult  female. 

Locality.  Canada:  British  Columbia,  Sumas.  30 
September  1895. 

Collector  A.  C.  Brooks.  Original  number  518. 
Condition.  Skin  and  skull  complete.  Mandible  dis- 
articulated. 

Type  Series.  3  paratypes;  B3700,  skin  and  skull, 
adult  female,  and  B5548,  skin  and  skull,  adult  male; 
B7435,  skull,  adult  male. 

Comments.  M.  m.  spissigrada  was  retained  as  a  val- 
id subspecies  by  Hall  (1981:  1022)  and  Wade- 
Smith  and  Verts  (1982:  1). 


Type  Specimens  of  Recent  Mammals  •  Hel^en  and  McFadden        155 


Genus  SPILOGALE  Gray,  1865 

Spilogale  ambarvalis  Bangs,  1 898b 

Proc.  Boston  Soc.  Nat.  Hist.,  28:  222,  15 

March. 

=  Spilogale  putorius  ambarvalis  Bangs, 

1898.  See  Van  Gelder  (1953:  255). 

Holotype.  B3481.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Florida,  Brevard  County, 

Oak  Lodge,  east  peninsula  opposite  Mieco.  30  Jan- 

uaiy  1895. 

Collector.  O.  Bangs.  Original  number  11. 

Condition.  Skin  and  skull  complete. 

Type  Series.  There  is  a  large  series  of  paratypes  in 

the  MCZ. 

Comments.  S.  p.  ambarvalis  was  retained  as  a  valid 

subspecies  by  Hall  (1981:  1014)  and  Kinlaw  (1995: 

1). 

Family  VIVERRIDAE  Gray,  1821 

Genus  EUPLERES  Doyere,  1835 

Eupleres  major  Layau6en,  1929 

Compt.  Rend.  Acad.  Sci.  Paris,  189:  198, 

22  July  22. 

=  Eupleres  goudotii  major  Lavauden, 

1929.  See  Albignac  (1973:  23). 

Syntypes.  MCZ  45691:  Skin,  skull,  and  postcranial 
skeleton.  Subadult  female,  MCZ  45962:  Skin,  skull, 
and  postcranial  skeleton.  Subadult  male. 
Locality.  Madagascar:  (Antsiranana),  foot  of  the 
Massif  Tsaratanna  (=Tsaratanana),  Upper  Sombi- 
rano  Valley,  above  village  of  Beangona,  1,500  m. 
April  1929. 
Collector  Lavauden. 

Condition.  MCZ  45691:  Skin  complete,  v^dth  bald 
spot  on  dorsum  and  tail  slightly  damaged.  Skull  and 
skeleton  complete.  Mandible  disarticulated.  Teeth 
removed  from  skull  but  present.  MCZ  45962:  Skin 
complete.  Skull  and  skeleton  complete.  Mandible 
disarticulated.  Teeth  removed  from  skull  but  pres- 
ent. 

Type  Series.  2  syntypes,  described  above. 
Comments.  These  are  the  two  specimens  from  G. 
Grandidier's  personal  collection  on  which  Lavau- 
den based  his  original  description  of  Enpleres  ma- 
jor Albignac  (1973:  23)  wrote  that  these  type  spec- 
imens were  "introuvable  [nowhere  to  be  found]." 

Family  HERPESTIDAE  Bonaparte,  1845 

Genus  GALIDICTIS  I.  Geoffrey,  1839 

Galidictis  grandidiensis  [sic]  Wozencraft, 

1986 

J.  Mammal.  67:  561,  8  August. 

=  Galidictis  grandidieri  Wozencraft,  1 986. 

See  Wozencraft  (1987:  198). 

Holotype.    MCZ  45983.   Skin,  skull,  and  postcranial 
skeleton.  Adult. 


Locality.  Madagascar  (no  further  data  available). 
The  locahty  of  the  paratype,  stored  in  the  AMNH, 
is  "Madagascar,  Lac  Tsimanampetsotsa,  24°08'  S, 
43°46'  E." 

Collector   No  collection  data  available.  The  holo- 
type is  part  of  the  collection  of  G.  Grandidier. 
Condition.    Skin   prepared  flat;   incomplete   (ven- 
trum  missing).  Skull  complete. 
Type  Series.  1  paraty^^e;  AMNH  100478,  sldn  and 
skull,  adult  male. 

Comments.  G.  grandidieri  was  considered  a  valid 
species  by  Wozencraft  (1993:  300)  and  Nowak 
(1999:  769). 

Family  FELIDAE  Fischer  de  Waldheim, 

1817 

Genus  LEPTAILURUS  Sevenzoy,  1858 

Fells  capensis  phillipsi  G.  M.  Allen,  1914d 

Bull.  Mus.  Comp.  Zool.,  58:  337,  July. 

=  Leptallurus  serval  phillipsi  (G.  M."  Allen, 

1914).  See  comments. 

Holotype.    MCZ    14908.    Skin    and    skeleton.    Adult 
male. 

Locality.  Sudan:  Blue  Nile,  El  Garef  10  Januaiy 
1913. 

Collector  J.  C.  PhiUips. 
Condition.  Skin  and  skull  complete. 
Type  Series.  Holotype  only. 

Comments.  G.  M.  Allen  (1939:  241)  used  the  name 
Felis  serval  phillipsi.  The  use  of  the  genus  Lep- 
tailunis  follows  Wozencraft  (1993:  292).  L.  s.  phil- 
lipsi was  retained  as  a  valid  subspecies  by  Smithers 
(1975:  7). 

Genus  LVA/XKerr,  1792 

Lynx  (Cervaria)  fasciatus  oculeus  Bangs, 

1899e 

Proc.  New  England  Zool.  Club,  1:  23,  31 

March. 

=  Lynx  rufus  callfornlcus  Mearns,  1897. 

See  Grinnell  and  DLxon  (1924:  346). 

Holotype.  B8633.  Skin  and  skull.  Adult  male. 

Locality.  (United  States):  Cahfomia,  Marin  Coun- 
ty, Nicasio.  11  December  1898. 
Collector  C.  A.  Allen.  Original  number  981. 
Condition.  Skin  and  skull  complete. 
Type   Series.    1   paratype;    B4789,   sldn  and  skull, 
adult  male. 

Lynx  gigas  Bangs,  1897e 

Proc.  Biol.  Soc.  Washington,  11:  50,  16 

March. 

=  Lynx  rufus  gigas  Bangs,  1897.  See 

Peterson  and  Downing  (1952:  11). 

Holotype.  B4951.  Skin  and  skull.  Adult  male. 

Locality.  Canada:  Nova  Scotia,  15  miles  (24.1  km) 
back  of  Bear  River.  11  December  1895. 


156         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Collector.  D.  R.  Ritchie. 
Condition.  Skin  and  skull  complete. 
Type  Series.  Holotype  only. 

Comments.  L.  r  gigas  was  retained  as  a  valid  sub- 
species by  Hall  (1981:  1054). 

Lynx  subsolanus  Bangs,  1897e 

Proc.  Biol.  Soc.  Washington,  11:  49,  16 

March. 

=  Lynx  canadensis  subsolanus  Bangs, 

1897.  See  ElHot  (1901:  296). 

Holotype.  B1190.  Sldn  and  skull.  Adult  male. 

Locality.  Canada:  Newfoundland,  Codroy.  13  June 
1894. 

Collector  E.  Doane. 
Condition.  Skin  and  skull  complete. 
Type  Series.  2  paratypes;  B5754,  skin  and  skull,  ju- 
venile female;  B3798,  skull,  adult  male. 
Comments.  L.  c.  subsolanus  was  retained  as  a  valid 
subspecies  by  Hall  (1981:  1051). 

Genus  PUMA  Jardine,  1834 

Felis  bangs!  Merriam,  1901 

Proc.  Washington  Acad.  Sci.,  3:  595,  11 

December. 

=  Puma  concolor  bangs!  {Memam,  1901). 

See  coinments. 

Holotype.  B8413.  Skin  and  skull.  Adult  male. 

Locality.  Colombia:  (La  Guajira),  Dibulla.  8  Oc- 
tober 1899. 

Collector.  W.  W.  Brown,  Jr. 
Condition.  Skin  and  skull  complete. 
Type  Series.   3  paratypes;   B8147,  skin  and  skull, 
adult  female,  and  "two  skulls  from  Peru,  in  the 
American  Museum  of  Natural  History." 
Comments.  Nelson  and  Goldman  (1929:  347)  used 
the  name  Felis  concolor  bangsi.  The  use  of  the  ge- 
nus  Puma    follows   Wozencraft    (1993:    296).    Re- 
tained as  a  valid  subspecies  by  Currier  (1983:  1). 

Fells  bangsi  costaricensis  Merriam,  1901 

Proc.  Washington  Acad.  Sci.,  3:  596,  11 

December. 

=  Puma  concolor  costaricensis  (Merriam, 

1901).  See  comments. 

Holotype.  MCZ  10118.  Skin  and  skull.  Adult  female. 
Locality.   Panama:  Chiriqui,  Boquete  (  =  Bajo  Bo- 
quete),  4,000  ft  (1,220  m).  22  April  1901. 
Collector  W.  W.  Brown,  Jr.  Original  number  337. 
Condition.  Skin  and  skull  complete. 
Type  Series.  There  is  a  series  of  paratypes  in  the 
MCZ. 

Comments.  Nelson  and  Goldman  (1929:  347)  used 
the  name  Felis  concolor  costaricensis.  The  use  of 
the  genus  Puma  follows  Wozencraft  (1993:  296). 
Retained  as  a  valid  subspecies  by  Currier  (1983:  1). 


Felis  coryi  Bangs,  1899b 

Proc.  Biol.  Soc.  Washington,  13:  15,  31 

Januaiy. 

=  Puma  concolor  coryi  {Bangs,  1899).  See 

comments. 

Holotype.  B7742.  Sldn  and  skull.  Adult  male. 

Locality.  (United  States):  Florida,  Brevard  County, 
"wildeniess  back  of  Sebastian".  1  Januaiy  1898 
Collector  F.  R.  Hunter. 
Condition.  Skin  and  skull  complete. 
Type    Series.    5   paratypes;    B5489,    adult    female, 
B5650,  adult  female,  B6992,  aduh  male,  B7743, 
adult   female,   B7744,  juvenile   female;   all  repre- 
sented by  sldn  and  skull. 

Comments.  Nelson  and  Goldman  (1929:  347)  used 
the  naine  Felis  concolor  coryi;  the  use  of  the  genus 
Puma  follows  Wozencraft  (1993:  296).  Felis  coryi 
Bangs,  1899  is  a  replacement  name  for  Felis  con- 
color floridana  Cory,  1896  (1896:  109).  Retained  as 
a  valid  subspecies  by  Currier  (1983:  1). 

Felis  improcera  Phillips,  1912 

Proc.  Biol.  Soc.  Washington,  25:  85,  4 

May. 

=  Puma  concolor  improcera  (Phillips, 

1912).  See  comments. 

Holotype.  MCZ  12704.  Skin  and  skull.  Adult  male. 
Locality.  (Mexico):  Lower  (  =  Baja)  California,  Cal- 
malli.  3  September  1911. 

Collector.  E.  W  Funcke.  Original  number  10. 
Condition.  Skin  and  skull  complete. 
Type  Series.  Holotype  only. 

Comments.  Nelson  and  Goldman  (1929:  347)  used 
the  name  Felis  concolor  improcera.  The  use  of  the 
genus  Puma  follows  Wozencraft  (1993:  296).  Re- 
tained as  a  valid  subspecies  by  Currier  (1983:  1). 

Order  CIMOLESTA  McKenna,  1975 

Family  MANIDAE  Gray,  1873 

Genus  PH/Ar/\G/A/aS  Rafinesque,  1821 

Phataginus  tricuspis  mabirae  G.  M.  Allen 

and  Loveridge,  1942 

Bull.  Mus.  Comp.  Zool,  89:  178, 

Februaiy. 

Holotype.    MCZ  39417.   Skin,  skull,  and  postcranial 
skeleton.  Adult  male. 

Locality.    Uganda:    (Buganda),    Chagwe,    Mabira 
Forest,  Mubango.  12  November  1938. 
Collector  A.  Loveridge. 

Condition.    Skin   and   skull   complete.    Postcranial 
skeleton  partial  (includes  atlas,  right  tibia,  and  right 
fibula). 
Type  Series.  Holotype  only. 


Type  Specimens  of  Recent  Mammals  •  Helgen  and  McFadden        157 


Comments.    Retained    as    a    valid    subspecies    by 
Meester  (1972:  2). 

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.    1896d.  On  a  small  collection  of  mammals 

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.   1896e.  The  cotton  mouse  Peroinyscus  gos- 

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.  1896g.  Preliminaiy  description  of  a  new  vole 

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.    1896h.    Some   new   mammals   from   Indian 


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.  1896i.  The  skunks  of  the  genus  Mephitis  of 

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.   1896J.  A  review  of  the  squirrels  of  eastern 

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.  1897a.  A  new  white-footed  mouse  from  Brit- 
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.  1897b.  Preliminar)'  description  of  the  New- 
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.  1897c.  Preliminary  description  of  a  new  race 


of  the  eastern  vole  from  Nova  Scotia.  American 
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.  1897d.  A  new  race  of  Gibb's  mole.  American 

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.  1897e.  Notes  on  the  lynxes  of  eastern  North 

America,  with  descriptions  of  two  new  species. 
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.   18971".   Description  of  a  new  red  fox  from 

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.   1897h.  On  a  small  collection  of  mammals 

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.  1897i.  A  new  race  of  pine  squirrel  from  the 

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.  1898a.  A  new  name  for  the  Nova  Scotia  fox. 

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.  1898c.  Descriptions  of  tvs'o  new  skunks  of  the 

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.    1898d.   Descriptions  of  the  Newfoundland 

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.   1898e.  The  eastern  races  of  the  American 

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.    1898f.    Description  of  a  new  white-footed 

mouse  from  the  Mount  Baker  Range,  British  Co- 
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.  1898g.  A  new  raccoon  from  Nassau  Island, 

Bahamas.  Proceedings  of  the  Biological  Society 
of  Washington,  12:  91-92. 

.  1898h.  Description  of  a  new  fox  from  Santa 

Marta,  Colombia.  Proceedings  of  the  Biological 
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.  1898i.  A  new  murine  opossum  from  Mar- 
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.   1898j.  A  list  of  the  mammals  of  Labrador. 

American  Naturafist,  32:  489-507. 

.  1898k.  Descriptions  of  some  new  mammals 

from  the  Sierra  Nevada  de  Santa  Marta,  Colom- 
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Washington,  12:  161-165. 

.    1898m.   A  new  race   of  the   little  hai"vest 


mouse  from  West  Virginia.  Proceedings  of  the 
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.  1898o.  On  Sciurus  variabilis  from  the  Santa 

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.  1898p.  A  new  rock  vole  from  Labrador.  Pro- 
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.    1898q.   A  new   Siginodon   from   the   Santa 

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.    1899a.  A  new  pigmy  Onjzomijs   from  the 

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.    1899b.  The  Florida  puma.   Proceedings  of 

the  Biological  Society  of  Washington,  13:  15-17. 

.   1899c.  A  new  race  of  striped  sperm ophile 

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.  1899d.  Notes  on  some  mammals  from  Black 

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.  1899e.  A  new  lynx  from  the  coast  of  Califor- 
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.  1899f.  A  new  race  of  chickaree.  Proceedings 

of  the  New  England  Zoological  Club,  1:  27-29. 

.  1899g.  Descriptions  of  two  new  pikas  from 

Western  North  America.  Proceedings  of  the  New 
England  Zoological  Club,  1:  39-42. 

.  1899h.  A  new  gray  fox  from  the  upper  Mis- 
sissippi Valley.  Proceedings  of  the  New  England 
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.  1899i.  Three  new  weasels  from  North  Amer- 
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.  1899j.  Descriptions  of  some  new  mammals 

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.  1899k.  A  new  bat  from  Colombia.  Proceed- 
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.    1900c.   Three   new  rodents  from   southern 

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.  1900d.  Description  of  a  new  squirrel  from 

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INDEX 


Type  Specimens  of  Recent  Mammals  •  Hel^en  and  McFadden        171 


abaconis 

Geocapromijs,  132 
abbotti 

Mijotis,  146 
abbreviata 

Neotonia,  121 
aberti 

Sciunis,  99 
abietorum 

Feromyscus,  125 
ablutus 

Leggada,  118 
acadicus 

Microtus,  115 

Neosorex,  138 

Zapus,  112 
aequalis 

Aedialops,  140 

Scalops,  140 

Scalopus,  140 
aestuans 

GueHinguetus,  106 

Sciiirus,  106 
Aethalops 

aequalis,  140 

alecto,  140 
Aethoglis 

argenteus,  129 

/jueij,  129 
Aethosciurus 

byatti,  106 

laetus,  106 
africanus 

Nycticeius,  147 
Afrosoricida,  101 
AgOMfi 

paca,  130 

virgatus,  130 
Agoutidae,  130 
akka 

FuniscUinis,  104 
Akodon 

apricus,  127 

teguina,  127 

xerampelinus,  127 
alacer 

Lepus,  134 

Sylvilagus,  134 
alashanicus 

Spermophilus,  107 
albicinctus 

Myotis,  146 
albigula 

Thyroptera,  147 
albiventer 

Nycticeius,  147 


Thyroptera,  147 
alboniger 

Hylopetes,  105 

Fteroinys,  105 
alecto 

Aethalops,  140 
alfari 

Microsciunis,  105 
alleni 

Dasymys,  118 

Heteromys,  112 

Hylomyscus,  118 

Lepus,  133 

Liomys,  112 

Macrotolagus ,  133 

Melomys,  118 

Rhinolophus,  143 

rt/ogfl 

Blarina,  137 
AZoKflrtfl 

luctuosa,  135 

palliata,  135 

pigra,  135 

trabeata,  135 
alpinus 

Sciuroptenis,  104 
altilis 

Scotophilus,  147 
amazonicus 

Nectomys,  121 
ambarvalis 

Spilogale,  155 
americana 

Martes,  152 
americanus 

Euarctos,  151 

Lept/s,  133,  134 

Odocoileus,  148 

Ursus,  151 
A77iefn(ia 

centurio,  143 

minor,  143 
ammodijtes 

Feromyscus,  125 
anastasae 

Feromyscus,  124,  125 

Scalops,  140 

Scalopus,  140 
anchietae 

Otomys,  120 
anchises 

Hylopetes,  105 

Fteromys,  105 
anetianus 

Fteropus,  141 
angolensis 

Mops,  145 


Antechinus 

mayeri,  100 

misim,  100 

noso,  100 
Antillogale 

marcanoi,  137 
aorensis 

Fteropus,  141 
Aplodontia,  103 

califomica,  103 

Columbiana,  103 

rainieri,  103 

7^(/a,  103 
Aplodontidae,  103 
Apodemus 

euxinus,  117 

mystacinus,  117 
apricus 

Akodon,  127 

Scotinomys,  127 
aquaticus 

Scalopus,  140 
aquilis 

Cerdocyon,  150 

Craseomys,  113 

Urocyon,  150 
aquilonius 

Fiber,  116 

Ondatra,  116 
araeum 

Flagiodonta,  132 
araneus 

Sorex,  138 
arcticus 

Hesperomys,  124 

Lepus,  133 

Rangifer,  149 
ArctogaZe 

longicauda,  153 

muricus,  153 

oribasus,  153 
Arcto??ii/.s 

avanis,  105 

flaviventer,  105 

igna-uus,  105 
arenarius 

Feromyscus,  126 
argentatus 

Feromyscus,  125 
argenteus 

Aethoglis,  129 

Graphiurus,  129 
arJeZ 

Fteropus,  141 
armatus 

Spermophilus,  107 
A/tiZjeus 


femurvillosum,  144 

lituratus,  144 

palmarum,  144 
Aitiodactyla,  145 
Arujco/a 

breweri,  114 

longipilis,  114 

riparia,  114 

rufidorsum,  114 

terraenovae,  114 
Arvicolinae,  112 
aslaeorientalis 

Neomeris,  148 

Neophocaena,  148 
Atalapha 

brachyotis,  146 

Suncus,  139 
atrata 

Martes,  152 

Mustela,  152 
atrodorsalis 

Thomomys,  111 
attioateri 

Feromyscus,  124 

Geogale,  101 
Triaenops,  143 
auritus 

Flecotus,  147 

Eothenomys,  113 

Microtus,  113 
austerulus 

Sigmodon,  127 
austini 

Fteropus,  141 
australis 

Cryptogale,  101 

Echymipera,  101 

Reithrodontomys,  127 
austrinus 

Geomys,  110 

Mannota,  105 
Arctomys,  105 

Cryptotis,  138 

Mephitis,  154 
bactrianus 

Mus,  121 
hadlus 

Ictidonujs,  108 

Spennophilus,  108 
hairdii 

Lepus,  133,  134 


172         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Mus,  98 

Peromyscii.s,  98 
haliolus 

Peromyscus,  126 
baluensis, 

Callosciunis,  103 
bangsi 

Fells,  156 

Glaucomys,  104 

Perognathus,  112 

Puma,  156 

Sciuroptenis,  104 

Viilpes,  151 
bangsii 

Lepus,  133 
barbouri 

Otomys,  121 
barrerae 

Octoniys,  130 

Tympanoctomys,  130 
Bathyergidae,  129 
hella 

Neotoma,  121 
belliis 

Peromyscus,  124 
Beluga 

concreta,  148 

declivis,  148 
beothucus 

Cants,  149 
bicolor 

Crocidura,  137 

Hipposideros,  142 

Oecomys,  122 
Blarina 

aloga,  137 

brevicauda,  137 

compacta,  137 
borealis 

hasiurus,  146 

Odocoileus,  148 

Peromyscus,  124 

Synaptomys,  116 

Tamias,  109 
Boromys 

torrei,  131 
borucae 

Sigmodon,  127 
Bovidae,  149 
brachyotis 

Atalapha,  146 

Lasiurus,  146 
brasiliensis 

Sylvilagus,  134 
brevicauda 

Blarina,  137 

Zygodontomys,  128 
brevicaudata 

Microgale,  102 

Arvicola,  114 


Microtus,  114 
brochus 

Syntheosciunis,  108 
browni 

Microsciunis,  105 

Sciurus,  105 
bninialis 

Mai-tes,  152 

Mustela,  152 
biifo 

Leggada,  118 

Mus,  118 
burrus 

Proechimys,  131 
byatti 

Aethosciurus,  106 

Paraxerus,  106 
caboti 

Rangifer,  149 
cacabatus 

Peromyscus,  124 

Castor,  109 
caecutiens 

Sorex,  138 
cfi/er 

Pedetes,  128 
californica 

Aplodontia,  103 
califoiiticus 

Dipodomys,  111 

Lynx,  165 

Scapanus,  140 
callida 

Dasyprocta,  130 
Callosciunis 

baluensis,  103 

erythraeus,  103 

ferrugineus,  103 

haemobaphes,  103 

medialis,  103 

primus,  103 
Caluromyidae,  99 
Caluromys 

cicur,  99 

lanatus,  99 
canadensis 

Castor  109 

Lontra,  152 

Lynx,  156 

Peromyscus,  125 
Canidae,  149 
canina 

Peropteryx,  142 
Cfinzs 

beothucus,  149 

familiaris,  150 

/tipus,  149,  150 
Cansumys 

canus,  117 
canus 


Cansumys,  117 
capensis 

Fells,  165 

Pedetes,  128 

Pipistrellus ,  146 
Capromyidae,  131 
Capromys 

nana,  132 

pilorides,  131 

relictus,  131 
capucinus 

Cebus,  136 
caraccioli 

Vampyrodes,  145 
Cariacus 

osceola,  148 
caribou 

Rangifer,  149 
carissinia 

My  Otis,  146 
Carnivora,  149 
carolinensis 

Sciurus,  106 
carpenteri 

Hijlobates,  136 
carroruni 

Oryzomys,  123 
cascadensis 

Lepus,  134 
castaneoventris 

Sciurus,  103 
castanonotus 

Sciurus,  99 
castanotus 

Sciurus,  99 
casta  nops 

Cratogeomys,  111 

Pappogeomys,  111 
Castor 

caecator,  109 

canadensis,  109 
Castoridae,  109 
cavator 

Macrogeomys,  110 

Orthogeomys,  110 
cayennensis 

Proechimys,  131 
Cebidae,  135 
Cebus 

capucinus,  136 

curtus,  136 
celatus 

Phenacomys,  116 
centu  rlo 

Ametrida,  143 
Cerdocyon 

aquilus,  150 

germanus,  150 

thous,  150 
Cervaria 

fasclatus,  155 


oculeus,  155 
Cei-vidae,  148 
Cetartiodactyla,  97 
Chaerophon 

leucostigma,  145 
Cheiromys 

laniger,  135 

madagascariensls,  135 
c/ierriei 

Zygodontomys,  128 
Chilonycteris 

pamellii,  143 

pusillus,  143 

torrei,  143 
Chionomys 

nivalis,  112 
chionopaes 

Dicrostonyx,  113 
chiriquensis 

Guerlinguetus,  106 

Sclunis,  106 
Chiroptera,  140 
Chlorotalpa 

tropicalis,  101 
chrotorrhinus 

Microtus,  114 
Chrysochloridae,  101 
Chnjsochloris 

stidilmanni,  101 

tropicalis,  101 
chrysogaster 

Lenimus,  114 
cicognonn 

Mustela,  152 
cicur 

Caluromys,  99 

Philander,  99 
Cimolesta,  156 
cinereoargenteus 

Urocyon,  150 
cJnereHS 

Sorgx,  139 
Cingulata,  101 
Citellus 

obscurus,  107 

.siccus,  107 
Clavlglls 

collaris,  129 

soleatus,  129 
clavlum 

Odocoileus,  149 
Clethrionomys 

gapperi,  113 

proteus,  113 
collaris 

Claviglis,  129 
colondylana 

Dasyprocta,  130 
colonus 

Geomys,  110 
coloratus 


Type  Specimens  of  Recent  Mammals  •  Hclfien  and  McFadden        173 


On/zoini/s,  123 
coliuiibiaiia 

Aplodoiifia,  103 
coliimbicnnis 

Geocapromijs,  132 
compacta 

Blarina,  137 
concoJor 

Felis,  156 

Puma,  156 
concreta 

Beluga,  148 
condylunts 

Mops,  145 
confucianus 

Niviventer,  119 
consobrinus 

Sciurofamias,  106 
cooper; 

Synaptovujs,  116 

Tainias,  109 
cory; 

Fe/;.s,  156 

Puma,  156 
Conjnorhinus 

phijllotis,  146 
casta  ricensis 

Felis,  156 

Pinna,  156 
cowani 

Microgale,  102 
Craseomijs 

aquilus,  113 
crassus 

Phenacornys,  116 
Cratogeomys 

castanops.  111 

rubellus.  111 
crgper 

ReUhrodontomys ,  127 
Cricetinae,  117 
Cricetomylnae,  117 
cricefulus 

Saccostomus,  117 
crinitus 

Peromyscus,  125 
Crocidura 

bicolor,  137 

fuscomurina,  137 

geata,  138 

hildegardeae,  138 

maurisca,  138 

phaios,  138 

tephragaster,  137 
cnisiiignim 

Pecan,  147 

Tayassu,  147 
Cryptogale 

a  u  straits,  101 
Cryptomys 

hottentotus,  129 


occlusus,  129 

ivhijtei,  129 
Cryptotis 

avia,  138 

tJioinasi,  138 
cubanus 

Geocaproinys,  132 

Solenodon,  137 
cumbedandius 

Geoim/s,  110 
c;/ppra 

Ochotona,  133 
ctirttis 

CeZpjf.s,  136 

Hipposideros,  142 
Damaliscus 

pJiiUipsi,  149 

pygargus,  149 
daphaenodon 

Sorex,  139 
darlingtoni 

Eptesicus,  145 

PipistreUus,  145 
Dasogale 

fontoynonti,  103 
Dasyinys 

alleni,  118 

inconitus,  118 
Dasypodidae,  101 
Dasyprocta 

callida.  130 

colombiana,  130 

leporina,  130 

noblei,  130 

nuchalis,  130 

punctata,  130 
Dasyproctidae,  130 
Dasypus 

hoplites,  101 

novemcinctus,  101 
Dasyuiidae,  100 
Das)airomoi"phia,  100 
Daubentonia 

n  ladagasca  riens  is ,  135 
Daubentoniidae,  135 
davidanus 

Sciurotamias,  106 
decaryi 

Microgale,  102 
decUvis 

Beluga,  148 
degener 

Lutra,  152 
delectonim 

Praomys,  119 
deletrix 

Vidpes,  150 
Delphinapterus 

leucas,  148 
Delphinus 

nielas,  148 


demidovii 

Galago,  135 
Dendromurinae,  117 
desmarestianus 

Heteromys,  111 

Oryzomys,  123 
diadeina 

Propithecus,  135 
Dicrostonyx 

chionopaes,  113 

exsid,  113 

groenlandicus,  113 

torquatus,  113 
Didelphidae,  100 
Didelphis 

marsupialis,  100 

particeps,  100 

pigra,  100 

virginiana,  100 
Didelphimorphia,  99 
Diplomys 

labilis,  131 
Dipodidae,  112 
Dipodomys 

californicus.  111 

ordii.  111 

pallidulus.  111 

palmeri.  111 
Dipodops 

ordii.  111 

palmeri.  111 
dispar 

Sorex,  139 
distincta 

Neotoma,  122 
dorcfls 

Damaliscus,  149 
dorsalis 

Tamias,  109 
dorsatum 

Erethizon,  130 
Dremomys 

flavior,  104 

pemyi,  104 

se?iex,  104 
droidtardi 

Microgale,  102 
douglasii 

Tamiasciunis,  109 
dupreanum 

Eidolon,  98 
Echimyldae,  131 
Echyniipera 

australis,  101 

kalabu,  100 

rufescens,  101 
Eidolon 

dupreanum,  98 

sakalava,  98 
eZegans 


Spennophilus,  108 
elongata 

Mephitis,  164 

Proct/on,  151 
Emballonuridae,  142 
energuinenos 

Mustela,  153 

Putorius,  153 

Microtus,  115 
Eonycteris 

glandifera,  140 

spelaea,  140 
Eothenomys 

aurora,  113 

eua,  113 

melanogaster,  113 

mucronatus,  113 
Epimys 

zappeyi,  119 
Eptesicus 

darlingtoni,  145 

pliasma,  146 

pumilus,  145 
erem/cus 

Hesperomys,  98 

Peromyscus,  98,  126 
Erethizon 

dorsatum,  130 

picinus,  130 
Erethizontidae,  130 
erigens 

Hipposideros,  142 
Erioryzomys 

monochromos,  128 

Mustela,  152,  153 
Erophylla 

sezekomi,  144 

syops,  144 
erythraeus 

Callosciurus,  103 
Euarctos 

americanus,  151 

sornhorgeri,  151 
Eulipotyphla,  97 
Eu pie  res 

goudotii,  155 

major,  155 
Eutamias 

minimus,  109 

neglectus,  109 
ewxmus 

Apod  emus,  117 
eufl 

Eothenomys,  113 
Evotomys 

proteus,  113 
exsputus 

Sigmodon,  127 


174         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


exsul 

Dicrostonyx,  113 
extinius 

Sciurus,  106 
familiaris 

Cants,  150 
fasciatus 

Cervaria,  155 

Lynx,  155 
fatuus 

Synaptomys,  116 
Felidae,  155 
Fells 

bangsi,  156 

capensis,  155 

concolor,  156 

coryi,  156 

costaricensis,  156 

floridana,  156 

improcera,  156 

philUpsi,  155 

serval,  155 
femurvillosu  m 

Artibeus,  144 
ferrugineus 

Callosciunis,  103 
feroidus 

Sigmodon,  128 
Fiber 

aquilonius,  116 

obscunis,  116 

rivalicius,  116 

zibethicus,  116 
fischeri 

Haplonycteris,  141 
flavicans 

Oecomys,  122 

Oryzomys,  122 
flavidus 

Isthniomys,  121 

Megadontomys,  121 
flavior 

Dremomys,  104 
flavipes 

Tat  era,  117 
flaviventer 

Arctomys,  105 
flaviventris 

Marmot  a,  105 
floridana 

Felis,  156 

Neotoma,  122 
floridanus 

Geomys,  110 

Sylvilagus,  134 
fontigenus 

Microtus,  115 
fontoynonti 

Dasogale,  103 
frenata 

Mustela,  152,  153,  154 


frenatus 

Putorius,  152 
fidvescens 

Oligoryzomys,  122 
fulviventer 

Mannosa,  99 
fumatus 

Myoniys,  119 

Praomys,  119 
Fimisciurus 

akka,  104 

pyrrhopiis,  104 

victoriae,  104 
furcuhis 

Triaenops,  143 
furvus 

Sigmodon,  128 

Urocyon,  150 
fuscipes 

Neotoma,  98 
fuscomurina 

Crocidura,  137 
fusus 

Peromyscus,  126 
Galago 

demidovii,  135 

orimis,  135 
Galagoides 

orinus,  135 
Galagonidae,  135 
Galidictis 

grandidiensis,  155 

grandidieri,  155 

Clethrionomys,  113 
geata 

Crocidura,  138 

Myosorex,  138 
Geocapromys 

abaconis,  132 

colutnbianus,  132 

cubanus,  132 

ingrahami,  132 

irrectus,  132 
GeogaZe 

aurita,  101 

orientalis,  101 
Geomyidae,  110 
Ggoinys 

austriniis,  110 

colonus,  110 

cumberlandius,  110 

floridanus,  110 

go#,  110 

pinetis,  110 

fwz.a,  110 
Gerbillinae,  117 
gerhillus 

Leggada,  119 
germanus 

Cerdocyon,  150 


gibbsi 

NeurotricJius,  140 
giganteiis 

Pteropus,  141 
gtgas 

Lynx,  155 
glaber 

Heterocephalus,  129 
glandifera 

Eonycteris,  140 
Glaucomys 

bangsi,  104 

lascivus,  104 

makkovikensis,  104 

querceti,  105 

sabrinus,  104 

uoZans,  104,  105 
Glossophaga 

longirostris,  144 
gloveralleni 

Procyon,  151 

Sorex,  138 
goffi 

Geomys,  110 
gorgonag 

Proechimys,  131 
gorilla 

Gorilla,  136 

Troglodytes,  136 
Gorilla 

gorilla,  136 
gossypinus 

Hesperomys,  123 

Peromyscus,  123,  124, 
125 
goudotii 

Eupleres,  155 
Grammonujs 

macmillani,  118 
granatensis 

Sciurus,  106,  107 
grandidiensis 

Galidictis,  155 
grandidieri 

Galidictis,  155 
grandis 

Orthogeonujs,  111 
Graphiurus 

argenteus,  129 

griseus,  129 

/lue^i  129 

lorraineus,  129 

microtis,  129 

murinus,  129 

schwabi,  129 

surdiis,  129 
griseus 

Graphiunis,  129 
griswoldi 

Tana,  134 
Guerlinmietus 


aestuans,  106 

chiriquensis,  106 
guyannensis 

Proechimys,  131 
gymnicus 

Sciurus,  109 

Tamiasciunis,  109 
haemobaphes 

Callosciunis,  103 

Sciurus,  103 
Haplonycteris 

fischeri,  141 
hardyi 

Zapus,  112 
haiyia 

Harpiocephalus,  146 
Hai~piocephalus 

harpia,  146 

nifuhis,  146 
harringtoni 

TateriUus,  117 
helleri 

Platyrrhinus,  144 
Herpestidae,  155 
hesperia 

Lonchophylla,  144 
Hesperomys 

arcticus,  124 

eremicus,  98 

gossypinus,  123 

leucopus,  124 

nebrascensis,  124 

sonoriensis,  124 
Heterocephalus 

glaber  129 

stygius,  129 
Heteromyidae,  111 
Heteromys 

alleni,  112 

desmarestianus.  111 

repens.  111 
hildegardeae 

Crocidura,  138 
Hipposideros 

bicolor,  142 

curtus,  142 

erigens,  142 

turpis,  143 
hirsutiis 

Sigmodon,  128 
hispid  a 

Suillomeles,  100 
hispidus 

Sigmodon,  127,  128 
Hominidae,  138 
hoplites 

Dasypus,  101 
hottentotus 

Cryptomys,  129 
hudsonica 

Lutra,  152 


Type  Specimens  of  Recent  Mammals  •  Helpen  and  McFadden        175 


luidsonlciis 

Sciunis,  109 

Tainiasciurus,  109 

Zapus,  112 
htieti 

Aedioglis,  129 

Graphiurus,  129 
huinidis 

Reifli  rodoiifoinys,  127 
hyacindiits 

Neiirotriclius,  140 
hijleae 

Proechimijs,  131 
Hylobates 

carpenteri,  136 

lai;  136 
Hylobatidae,  136 
Hijlomyscus 

alleni,  118 

simus,  118 
Hijlopetes 

alboniger,  105 

anchises,  105 

oriniis,  105 

phaijrei,  105 
hypoxanthus 

Oenomys,  119 
Hypudaeus 

nivicola,  112 
Ictidomys 

badius,  108 

trideceinllneatus,  108 
Idionycteris 

phyllotis,  146 
ignaua 

Marmot  a,  105 
jgnfluxs 

Arctomys,  105 
igni/er 

Rhinolophus,  143 
illectiis 

Oecotm/s,  122 

Oitjzomys,  122 
impiger 

Reldirodontoinys,  127 
improcera 

Fells,  156 

Puma,  156 
incitatus 

Lepus,  134 

Sylvilagus,  134 

Tapeti,  134 

Dasymys,  118 
Indridae,  135 
ingrahami 

Geocapromys,  132 
innultus 

Mictomys,  116 

Synaptomys,  116 
insulanus 


Peromyscus,  125 
insularis 

Pteropus,  98 

Geocapromys,  132 
irroratus 

Liomys,  112 
Isdimomys 

flavidus,  121 
kalabit 

Echymipera,  100 

T«f<?ra,  117 
kivtiensis 

Thamnomys,  120 
kodiacensis 

Spennopliilus,  99 

Microtus,  115 

Sorex,  138 
labilis 

Diplomys,  131 

Loncheres,  131 
lacustris 

Otomxjs,  120 
ladas 

Zapus,  112 
laetus 

Aethosciunis,  106 
Lagomoi'plia,  133 
lambertoni 

Nesomys,  120 
lanatus 

Cahiromys,  99 
laniger 

Cheiromys,  135 

Pteropus,  98 
lanigera 

Pteropus,  98 

Hylobates,  136 
lascivus 

Glaucomys,  104 

Sciuroptenis,  104 
Lasiurus 

borealis,  146 

brachyotis,  146 
latimanus 

Scapanus,  140 
laxatina 

Lontra,  142 
lecontii 

Reithrodontomys,  127 
Leggada 

ablutus,  118 

fou/o,  118 

gerhtllus,  119 
Lemr7ius 

chrysogaster,  114 

paulus,  114 

sibiricus,  114 


lepida 

Neotoma,  121 
Leporidae,  143 
leporina 

Dasyprocta,  130 
Leptailunis 

phillipsi,  155 

serval,  155 
LepM5 

alacer,  134 

alleni,  133 

americanus,  133,  134 

arcticus,  133 

Z?air<ii,  133,  134 

bangsii,  133 

cascadensis,  134 

incitatus,  134 

palitans,  133 

paludicola,  134 

stnithopus,  133 

sylvaticus,  134 

transltionalis,  134 

Delphinaptenis,  148 
leucogaster 

Scotophilus,  147 
leucopus 

Hesperomys,  124 

Peromyscus,  126 
leucostigma 

Chaerophon,  145 

Mops,  145 
levipes 

Melomys,  118 
hiomys 

alleni,  112 

irroratus,  112 
Lipotyphla,  136 
lituratus 

Artibeus,  144 
Loncheres 

labilis,  131 
Lonchophylla 

hesperia,  144 
longicauda 

Arctogale,  153 

Putorius,  153 
longicaudata 

Microgale,  102 
longimembis 

Perognathus,  112 
longipilis 

Arvicola,  114 
longirostris 

Glossophaga,  144 
Lonfra 

canadensis,  152 

laxatina,  152 

Sciurus,  109 
Tamiasciurus,  109 


lorraineus 

Graphiurus,  129 
Zotor 

Procyon,  151 
louisianae 

Odocoileus,  148 
lucifugus 

My  Otis,  146 
luctuosa 

Alouatta,  135 
ludovicianus 

Sciurus,  106 
lupus 

Canis,  149,  150 
lutensis 

Lutreola,  153 

Mustela,  153 

Putorius,  153 

degener,  151 

hudsonica,  151 

uago,  151 
Lutreola 

lutensis,  153 

vulgivagus,  154 
Lynx 

calif ornicus,  155 

canadensis,  155,  156 

fasciatus,  155 

gigas,  155 

oculeus,  155 

nifus,  155 

subsolanus,  156 
mabirae 

Phataglnus,  156 
macmillani 

Grammomys,  118 
Macrogeomys 

cavator,  110 

pansa,  110 
macropygmaeus 

Sorex,  138 
macrotis 

Nycteris,  142 

Peropteryx,  142 
Macrotolagus 

alleni,  133 

palitans,  133 
macrou  ra 

Odocoileus,  148 
macnirus 

Sorex,  139 
madagascariensis 

Cheiromys,  135 

Daubentonia,  135 

Nycteris,  142 

Rousettus,  141 
major 

Eupleres,  155 

Vampyrodes,  145 
makkovikensis 


176         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Glaucomys,  104 

Sciuroptenis,  104 
maniculatus 

Peromyscus,    98,    124, 
125 
Manidae,  156 
marcanoi 

Antillogale,  137 

Solenodon,  137 
Marmosa 

fulviventer,  99 

mitts,  99 

robinsoni,  99,  100 
Marmosidae,  99 
Marmota 

avara,  105 

flaviventris,  105 

ignava,  105 

monax,  105 
inarsupialis 

Didelphis,  100 
Martes 

americana,  152 

atrata,  152 

bnimalis,  152 
maurisca 

Crocidura,  138 
mayeri 

Antechinus,  100 
rnaynardi 

Procyon,  151 
meamsz 

Saccostomus,  117 
niedialis 

Callosciunis,  103 
niedioximii.s 

Mictomys,  116 

Synaptomys,  116 
Megadontomys 

flavidus,  121 
megaphyllus 

Rhinolophus,  143 
melanogaster 

Eothenomys,  113 
melanops 

Taterillus,  117 
melanotis 

Oryzomys,  123 
melanotus 

Praomys,  119 
meZas 

Delphinus,  148 

Myoictis,  100 
Melornys 

alleni,  118 

levipes,  118 

mollis,  118 

moncktoni,  118 

ruhex,  118 

stevensi,  118 
mephitica 


Mephitis,  154 
Mephitidae,  154 
mephitis 

Mephitis,  154 
Mephitis 

avia,  154 

elongata,  154 

mephitica,  154 

mephitis,  154 

mesomelas,  154 

scrutator,  154 

spissigrada,  154 
Mesocap  romys 

nanus,  132 
mesomelas 

Mephitis,  154 

Neotoma,  122 
mexicanus 

Peromyscus,  124 
Microgale 

brevicaudata,  102 

cowani,  102 

decanji,  102 

droiihardi,  102 

longicaiidata,  102 

parvula,  102 

principtda,  102 

prolixicaudata,  102 

pu/Zfl,  102 
Microsciurus 

alfan,  105 

browni,  105 
microtis 

Graphiuriis,  129 
Microtus 

acadtcus,  115 

aurora,  113 

breweri,  114 

chrotorrhinus,  1 14 

enixus,  115 

fontigenus,  115 

koreni,  115 

mucronatus,  113 

oeconomus,  115 

pennsylvanicus,  114, 
115 

provectus,  115 

ravus,  114 

shattucki,  115 

terraenovae,  114 

Nesophontes,  136 
Mictomys 

innuitus,  116 

medioximus,  116 
minimus 

Tamias,  99,  109 

Ametrida,  143 
Suncus,  139 


minusculus 

Scapanus,  140 
mjsczx 

Sorex,  139 
misim 

Antechinus,  100 
mitis 

Marmosa,  99 
mollipilosus 

Tamiasciunis,  109 
mollis 

Melonujs,  118 

Stenomys,  120 
Molossidae,  145 
monax 

Marmota,  105 
moncktoni 

Melomys,  118 
monochromos 

Erioryzomys,  128 

Oryzomys,  128 

Thomasomys,  128 
Monodontidae,  148 
monoensis 

Pteropus,  141 
Mops 

angolensis,  145 

condylunis,  145 

leucostigma,  145 

orientis,  145 
Mormoopidae,  143 
mortigena 

Mustela,  152 
morulus 

Sciurus,  107 
mucronatus 

Microtus,  113 
munda 

Mustela,  154 
mundus 

Putorius,  154 
muricola 

Myotis,  146 
initnctis 

Arctogale,  153 

Mustela,  153 

Putorius,  153 
Muridae,  112 
Murinae,  117 
murinus 

Graphiuriis,  129 

Pseudohydromys,  120 
Mms 

bactrianus,  119 

bairdii,  98 

Z;r//o,  118 

niusculus,  119 

tantillus,  119 

tenellus,  119 
musculiis 

Mus,  119 


Mustela 

atrata,  152 

bnimalis,  152 

cicognanii,  152 

energiimenos,  153 

erminea,  152,  153 

frenata,  152,  153,  154 

lutensis,  153 

mortigena,  152 

munda,  154 

muricus,  153 

neomexicana,  152 

nivalis,  153 

noveboracensis,  153 

occisor,  153 

oribasus,  153 

richardsonii,  152 

rixosa,  153 

vison,  153,  154 

vulgivaga,  154 
Mustelidae,  152 
Myoictis 

nielas,  100 

wallacei,  100 

wavicus,  100 
Myomys 

funmtus,  119 
Myopus 

schisticolor,  115 

thayeri,  115 
Mi/osorex 

geata,  138 
Myofjs 

abbotti,  146 

albicinctus,  146 

carissima,  146 

lucifugus,  146 

muricola,  146 

nugax,  146 

sodalis,  146 
Myoxidae,  129 
mystacinus 

Apodemus,  117 
nana 

Capromys,  132 

Nycteris,  142 
nanus 

Mesocapromys,  132 
nrtso 

Antechinus,  100 

Rhynchiscus,  142 
natator 

Oryzomi/s.  123 
naiyws 

Oryzomi/s,  122 
nebrascensts 

Hesperomys,  124 

Peromyscus,  124 
Nectomys 

aniazonicus,  121 

squamipes,  121 


Type  Specimens  of  Recent  Mammals  •  Helgen  and  McFadden        111 


neglectus 

Tainias,  109 
Neomeris 

asiaeorientalis,  148 
neomexicana 

Musfelfi,  152 
neoinexica)}its 

Putoriiis,  152 
Neophocaena 

asiaorientalis,  148 

phocaenoides,  148 
Neosoi-ex 

acadicus,  138 

palttstris,  138 
Neotonia 

abhreviata,  121 

beUa,  121 

distincta,  122 
floridana,  122 

fuscipes,  98 

lepida,  121 

mexicana,  122 

riibida,  122 
nesaeiis 

Scitinis,  107 
Nesomylnae,  120 
Nesomys 

lambertoni,  120 
Nesophontes 

micriis,  136 
Nesophontidae,  136 
Neil  rot richits 

gibbsi,  140 

Jiyaciiidihiiis,  140 

Sciiiriis,  106 
nigrictihis 

Peromysciis,  125 

StenoDiys,  120 
nitellinits 

Nyctoinys,  122 
nivalis 

Chionomys,  112 

Mustela,  153 
nivicola 

Hypudaeiis,  112 
Niviventer 

confuciainis,  119 
noblei 

Dasyprocta,  130 

Pipistrelliis,  147 

Putorius,  153 

Scabrifer,  147 
noveboracensis 

Mustela,  153 

Putorius,  153 
novemcinctus 

Dasypus,  101 
nuchalis 


Dasyprocta,  130 
nugax 

My  Otis,  146 

Steatomys,  117 
Nycteridae,  142 
Nycteris 

macrotis,  142 

madagascariensis,  142 

nana,  142 

revoili,  97 

tristis,  142 
Nycticeiiis 

africanus,  147 

albiventer,  147 

schlieffeni,  147 
Nyctomys 

nitellinus,  122 

sumichrasti,  122 
ohsciinis 

Citellus,  107 

FAer,  115 

Ondatra,  115 
obsoletus 

Spennophilus,  99 
occidentalis 

Paramicrogale,  102 
occisor 

Mustela,  153 

Putorius,  153 
occlusus 

Cryptonu/s,  129 
Ochotona 

cupppes,  133 

princeps,  133 

saxatilis,  133 
Ochotonidae,  133 
ochraceiis 

Thamnomys,  118 
Octodontidae,  130 
Octomys 

barrerae,  130 

Cervaria,  155 

Lynx,  155 
ocythoiis 

Urocyon,  150 
Odocoileus 

americanus,  148 

borealis,  148 

clavium,  149 

louisianae,  146 

macroura,  146 

osceola,  146 

virginianus,  146,  147 

bicolor,  122 
flavicans,  122 
illectus,  122 
trabeatus,  122 
oecono»?i;<s 


Microtus,  115 

Oenomys 

hypoxanthus,  119 
talangae,  119 

Oligoryzornys 
fulvescens,  122 
vegetus,  123 

Ondatra 

ac/uilonius,  116 
obscunis,  116 
rivalicius,  116 
zibethictis,  116 

orariiis 

Sciunis,  109 
Zapus,  112 

ordii 

Dipodomys,  111 
Dipodops,  111 

oreas 

Peromyscus,  126 

oribasus 

Arctogale,  153 
Mustela,  153 
Putorius,  153 

orientalis 

Geogale,  101 

orientis 
Mops,  145 

onn«.S' 

Gala  go,  135 
Galagoides,  135 
Hylopetes,  105 
Pteromys,  105 

Orthogeoniys 
cavator,  110 
grandis.  111 
pansa,  110 
p/((fo.  111 

Oryzoinys 

carrorum,  123 
coloratus,  123 
devius,  123 
flavicans,  122 
fulvescens,  122 
illectus,  122 
nielanotis,  123 
monochronios,  128 
natator,  123 
navus,  122 
palustris,  123 
rostratus,  123 
vegetus,  123 

osceoZa 

Cariacus,  148 
Odocoileus,  146 

Otomops 

papuensis,  145 

Otomyinae,  120 

anchietae,  120 
barboiiri,  121 


lacustris,  120 
typus,  121 
uzungwensis,  121 

Praoinys,  119 

Agouti,  130 
Pachycyon 

robustus,  150 
paitana 

Tupaia,  134 
palitans 

Lepus,  133 

Macrotolagus,  133 
palliata 

Alouatta,  135 
pallidulus 

Dipodomys,  111 
pallidus 

Spennophilus,  99 

Tainias,  99 
pahnarius 

Peromyscus,  125 
pabnarum 

Artibeus,  144 
pahneri 

Dipodomys,  111 

Dipodops,  111 
paludicola 

Lepus,  134 

St/lvilagus,  134 
palustris 

Neosorex,  138 

Onjzomys,  123 

Sorex,  138 

Sylvilagus,  134 
pansa 

Macrogeomys,  1 10 

Ortliogeonujs,  110 
Pappogeomys 

castanops.  111 

ntbellus.  111 
papuensis 

Otomops,  145 
Paracaproniys,  132 
Paramicrogale 

occidentalis,  102 
Paraxenis 

byatti,  106 

vexillarius,  106 
paniellii 

Chilonycteris,  143 

Pteronotus,  143 
parryi 

Spennophilus,  99 
particeps 

Didelphis,  100 
parvula 

Microgale,  102 
paulus 

Leninuis,  114 


178         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  2 


Pecari 

ci-usnigrum,  147 

tajacu,  147,  148 

torvus,  148 
Pedetes 

cafer,  128 

capensis,  128 

taborae,  128 
Pedetidae,  128 
pennsylvanica 

Vulpes,  150,  151 
pennsylvanicus 

Microtus,  114,  115 
Peramelia,  100 
pernyi 

Dremomys,  104 
Perognathus 

hangsi,  112 

longiinembris,  112 
Peromyscus 

abietoruin,  125 

ammodytes,  125 

anastasae,  124,  125 

arenarius,  126 

argentatus,  125 

attwateri,  124 

bairdii,  98 

baliolus,  126 

foeZZtis,  124 

borealis,  124 

cacabatiis,  124 

canadensis,  125 

crinitus,  125 

eremicus,  98,  126 

fusus,  126 

gossypinus,  123,  124, 
125 

insidanus,  125 

leucopus,  126 

maniculatus,  98,  124, 
125 

mexicaniis,  124 

nebrascensis,  124 

nigriculus,  125 

oreas,  126 

pahnarius,  125 

phasma,  126 

polionotus,  126 

rhoadsi,  126 

saturatus,  126 

scitulus,  125 

subgrisetis,  126 

texanus,  126 
Peropteryx 

canina,  142 

macrotis,  142 

phaea,  142 
Peroiyctidae,  100 
perneri 

Propithecus,  135 
personatus 


Sorex,  139 
phaea 

Peropteryx,  142 
phaios 

Crocidura,  138 
phasma 

Eptesicus,  147 

Peromyscus,  126 

Pipistrellus,  147 
Phataginus 

niabirae,  156 

tricuspis,  156 
phayrei 

Hylopetes,  105 

Pterornys,  105 
Phenacomys 

celatus,  116 

crassus,  116 

ungava,  116 
Philander 

ciciir,  99 
philippinensis 

Rhinolophus,  143 
phillipsi 

Damaliscus,  149 

Fe/is,  155 

Leptatlunis,  155 
phocaenoides 

Neophocaena,  148 
Phocoenidae,  148 
Phyllostomidae,  143 
phyllotis 

Corynorhinus,  146 

Idionycteris,  146 
picnii/s 

Erethizon,  130 
pigra 

Alouatta,  135 

Didelphis,  100 
piloroides 

Capromys,  131 
pinetis 

Geomys,  110 
Pipistrellus 

capensis,  147 

notius,  147 

phasma,  147 

rendalli,  147 
Plagiodonta 

araeum,  132 
Platyrrhinus 

helleri,  144 

umbratus,  144 
Plecotus 

aiiritus,  147 

sacrimontis,  147 
pluto 

Orthogeomys,  111 
poeyanus 

Solenodon,  137 
polionotus 


Peromyscus,  126 
Praomys 

delectorum,  119 
fumatus,  119 

melanotus,  119 

oweni,  119 

tuHbergi,  119 
pratensis 

Steatomys,  117 
Primates,  135 
primus 

Callosciimts,  103 
pnuceps 

Ochotona,  133 
principula 

Micro  gale,  102 
prisons 

Rhynchiscus,  142 
Procyon 

elucus,  151 

gloveralleni,  151 

Zofor,  151 

maynardi,  151 
Procyonidae,  151 
Proechimys 

burnis,  131 

cayennensis,  131 

gorgonae,  131 

guyannensis,  131 

hyleae,  131 

semispinosus,  131 
p  rolixicaudata 

Microgale,  102 
Propithecus 

diadema,  135 

perrieri,  135 
proteus 

Clethrionomys,  113 

Evotomys,  113 
provectus 

Microtus,  115 
Pseudohydromys 

murinus,  120 
Pferojnys 

alboniger,  105 

anchises,  105 

orinus,  105 

phayrei,  105 
Pteronotus 

pamellii,  143 

pusillus,  143 

quadridens,  143 
Pteropodidae,  140 
Pteropus 

anetianus,  141 

aorensis,  141 

rtrie/,  141 

austini,  141 

giganteus,  141 

insidaris,  98 

laniger,  98 


lanigera,  98 

monoensis,  141 

rayneri,  141 

woodfordi,  141 
pulla 

Microgale,  102 
Pu??ifl 

hangsi,  156 

concolor,  156 

coryi,  156 

costaricensis,  156 

improcera,  156 
punctata 

Dasyprocta,  130 
pusillus 

Chilonycteris,  143 

Pteronotus,  143 
putorius 

Spilogale,  155 
Putorius 

energumenos,  153 
frenatus,  152 

longicauda,  153 

lutensis,  153 

mundus,  154 

muricus,  153 

neomexicanus,  153 

notius,  153 

noveboracensis,  153 

occisor,  153 

oribasus,  153 

rixosus,  153 

vison,  153 

vidgivagus,  154 

xanthogenys,  154 
pj/gargMS 

Damaliscus,  149 
pyrrhopus 

Funisciunis,  104 
quadridens 

Pteronotus,  143 
qiiadrivittatus 

Tamias,  99,  109 
querceti 

Glaucomys,  105 

Sciuropterus,  105 
ramieri 

Aplodontia,  103 
Rangifer 

arcticus,  149 

caboti,  149 

caribou,  149 

tarandus,  149 

terraenovae,  149 
rat>u.s 

Microtus,  114 
rayneri 

Pteropus,  141 
Reithrodontomys 

australis,  127 

creper,  127 


Type  Specimens  of  Recent  Mammals  •  Helaen  and  McFadden        179 


hinutills,  127 

itnpiger,  127 

lecontii,  127 

sumichrasti,  127 

viilcaniiis,  127 
relicttis 

Capromys,  131 
rendalli 

Pipistrelhis,  147 
repens 

Heteroimjs,  111 

Nijcteris,  97 
Rhinolophidae,  142 
Rliinolopliiis 

alleni,  143 

ig;nifer,  143 

megaphyllus,  143 

philippinensis,  143 
rhoadsi 

Peroinijscus,  126 
Rhynchiscus 

naso,  142 

priscus,  142 
RJn/nchonycteris 

naso,  142 
richardsonii 

Mustela,  152 
riparia 

Arvicola,  114 
rivalicius 

Fiber,  116 

Ondatra,  116 
rixosrt 

Mustela,  153 
nxosrts 

Piitorius,  153 
robinsoni 

Mannosa,  99,  100 
roboratus 

Sorex,  139 
robustiis 

PacJu/cyon,  150 
Rodentia,"  103 
ro  stratus 

Oryzomys,  123 
Rousettus 

madagasca  riens  is ,  141 
nibellus 

Cratogeomys,  111 

Pappogeoniys,  111 
nibex 

Melomys,  118 
rubida 

Neotoma,  122 
rubricatus 

Vulpes,  151 
nibricosa 

Vulpes,  150,  151 

Aplodontia,  103 


nifescens 

Echymipera,  101 
rufidorsunx 

Arvicola,  114 
rufulus 

Haiyioceplialus,  145 
rufus 

Lynx.  155 
sabrinus 

Glaucoinys,  104 

Sciuroptenis,  104 
Saccostomus 

cricetulus,  117 

meamsi,  117 
sacrimontis 

Plecotus,  147 
sakalava 

Eidolon,  98 
saltuensis 

Sciunis,  107 
sanctaeniartae 

Sigmodon,  128 
sanguinidens 

Sorex,  139 
saturatus 

Peroniyscus,  126 
saxatilis 

Ochotona,  133 
Scabrifer 

notius,  147 
Scalops 

aereus,  140 

anastasae,  140 

texanus,  140 
Scalopus 

aereus,  140 

anastasae,  140 

aquaticus,  140 
Scandentia,  134 
Scapanus 

califomicus,  140 

latirnanus,  140 

minusculus,  140 
schisticolor 

Myopus,  115 
schliejfeni 

Nycticeius,  147 
Schwab  i 

Graphiurus,  129 
scitulus 

Peroniyscus,  125 
Sciuridae,  103 
Sciuroptenis 

alpinus,  104 

bangsi,  104 

lascivus,  104 

niakkovikensis,  104 

querceti,  105 

sabrinus,  104 

.siZiis,  104 

ooZons,  104,  105 


Sciurotaniias 

consobrinus,  106 

davidianus,  106 

thayeri,  106 
Scit<n<s 

aberti,  99 

aestuans,  106 

browni,  105 

carolinensis,  106 

castaneoventris,  103 

castanonotus,  99 

castanotus,  99 

chiriquensis,  106 

extimus,  106 

granatensis,  106,  107 

gymnicus,  109 

haemobaphes,  103 

hudsonicus,  109 

loquax,  109 

ludovicianus,  106 

mondus,  107 

nesaeus,  107 

niger,  106 

orarius,  109 

saltuensis,  107 

variabilis,  107 

vicinus,  106 

vulpinus,  106 

ijucatanensis,  106 
Scotinomys 

apricus,  127 

teguina,  127 

xeranipelinus,  127 
Scotophilus 

altilis,  147 

leucogaster,  147 
scrutator 

Mephitis,  154 
seniispinosus 

Proechimys,  131 
senex 

Drenionujs,  104 

Zygodontomys,  128 
serval 

Felis,  155 

Leptailunis,  155 
Setifer 

setosus,  103 
setosus 

Setifer,  103 

Erophylla,  144 
shattucki 

Microtus,  115 
sibiricus 

Lemrnus,  114 
siccus 

Citellus,  107 
Sigmodon 

austendus,  127 


borucae,  127 
exsputus,  127 
fervidus,  128 
furvus,  128 
hirsutus,  128 
hispidus,  127,  128 
sanctaeniartae,  128 
spadicipygus,  128 

Sigmodoiitinae,  121 

siZus 

Sciuroptenis,  104 

SiHUiS 

Hylomyscus,  118 
sodalis 

Myotis,  146 
soleatus 

Claviglis,  129 
Solenodon 

cubanus,  137 

marcanoi,  137 

poeyanus,  137 
Solenodontidae,  137 
sonoriensis 

Hesperomys,  124 
Sorex 

araneus,  138 

caecutiens,  138 

cinereus,  139 

daphaenodon,  139 

dispar,  139 

gloveralleni,  138 

koreni,  138 

inacropygniaeus,  138 

niaci~urus,  139 

miscix,  139 

palustris,  138 

personatus,  139 

roboratus,  139 

sanguinidens,  139 

tundrensis,  138 

ultinuis,  138 

oir,  139 
Soricidae,  137 
sornborgeri 

Euarctos,  151 

L/rs((s,  151 
soro/- 

Trtfera,  117 
spadicipygus 

Sigmodon,  128 
spelaea 

Eonycteris,  140 
Spennophilus 

alashanicus,  107 

arniatus,  107 

badius,  108 

elegans,  108 

kodiacensis,  99 

obsoletus,  99 

pallidus,  99 

parryi,  99 


180         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No. 


spilosoma,  99 

texensis,  108 

tridecemlineatus,  99, 
108 
Spilogale 

ainbarvalis,  155 

putorius,  155 
spissigrada 

Mephitis,  154 
squarnipes 

Nectomys,  121 
Steatomys 

nyasae,  117 

pratensis,  117 
Stenoinys 

mollis,  120 

niobe,  120 

stevensi,  120 

verecundus,  120 

Melomys,  118 

Stenoinijs,  120 
striatus 

Tamias,  107 
stnithopus 

Lepiis,  133 
stuhhnanni 

Chnjsochloris,  101 
stygius 

Heterocephalus,  129 
subgriseus 

Peromyscus,  126 
suhsolanus 

Lynx,  156 
Suillomeles 

hispida,  100 
sumichrasti 

Nyctomys,  122 

Reithrodontomys,  127 
Suncus 

ater,  139 

minor,  139 

varilla,  139 

Graphiunis,  129 
sylvaticus 

Lepus,  134 
Sylvilagus 

alacer,  134 

brasiliensis,  134 
floridanus,  134 

incitatus,  134 

paludicola,  134 

palustris,  134 

transitionalis ,  134 
Synaptomys 

boreal  is,  116 

cooperi,  116 
fatuus,  116 

innuitus,  116 

medioxitnus,  116 


Syntheosciwins 

broclius,  108 
syops 

Erophylla,  144 
taborae 

Pedetes,  128 
tajacu 

Pecari,  147,  148 
talangae 

Oenomys,  119 
Talpidae,  140 
Tamias 

borealis,  109 

cooperi,  108 

dorsalis,  108 

minim.us,  99,  109 

neglectus,  109 

pallidus,  99 

quadrivittatus,  99 

striatus,  109 

townsendii,  108 

venustus,  109 
Tamiasciunis 

douglasii,  109 

gym,nicus,  109 

hudsonicus,  109 

loquax,  109 

mollipilosus,  109 
Tanfl 

grisiooldi,  134 

tona,  134 
fana 

Tana,  134 

Tupaia,  134 
tantilliis 

Mas,  119 

incitatus,  134 
ta  rand  us 

Rangife7',  149 
Tafera 

flavipes,  117 

kempi,  117 

soror,  117 

valida,  117 
Taterillus 

liarringtoni.  117 

melanops,  117 
Tayassu 

crusnignim,  147 

torvus,  148 
Tayassuidae,  147 
teguina 

Akodon,  127 

Scotinomys,  127 
tenellus 

Mus,  119 
Tenrecidae,  101 
tephragaster 

Crocidura,  137 
terraenovae 


A)~vicola,  114 

Microtus,  114 

Rangifer,  149 
texanus 

Peromyscus,  126 

Scalops,  140 
fexe'n.sz.s 

Spernwpliilus,  108 
Thamnomys 

kivuensis,  120 

ochraceus,  118 

venustus,  120 
thayeri 

My  opus,  115 

Sciurotamias,  106 
thoniasi 

Crypt  Otis,  138 
Thomasomys 

nionochromos,  128 
Thoniomys 

atrodorsalis.  111 

unibrinus.  111 

Cerdocyon,  150 
Thyroptera 

albigula,  147 

alhiventer,  147 

tricolor,  147 
Thyropteridae,  147 
torquatus 

Dicrostonyx,  113 
torrei 

Boromys,  131 

Chilonycteris,  143 
toruw5 

Pecari,  148 

Tayassu,  148 
townsendii 

Tamias,  109 
trabeatiis 

Alouatta,  135 

Oecomys,  122 
transitionalis 

Lepus,  134 

Sylvilagus,  134 
Triaenops 

aurita,  143 
furculus,  143 
fricoZor 

Thyroptera,  147 
tricuspis 

Phataginus,  156 
tridecemlineatus 

Spennophilus,  99,  108 
trinotatus 

Zapus,  112 

Nycteris,  142 
Troglodi/tes 

gorilla,  136 
tropicalis 


Chlorotalpa,  101 

Chnjsochloris,  101 
tullbergi 

Praomys,  119 
tundrensis 

Sorex,  138 
Tupaia 

paitana,  134 

tona,  134 
Tupaiidae,  134 
turpis 

Hipposideros,  143 

Geo  my  s,  110 
Tympanoctomys 

barrerae,  130 
typus 

Otomys,  121 
ultimus 

Sorex,  138 
umbratus 

Platyrrhinus,  144 

Vampyrops,  144 
umbrinus 

Thomonu/s,  111 
ungava 

Phenacomys,  116 
Urocyon 

aquilus,  150 

cinereoargenteus,  150 
furvus,  150 

ocythous,  150 
Ursidae,  151 

americanus,  151 

somborgeri,  151 
uzungwensis 

Otomys,  121 
vafra 

Vulpes,  150,  151 
uaga 

L«fra,  152 
valida 

Tatera,  117 
Vampijrodes 

caraccioli,  145 

major,  145 
Vampyrops 

umbratus,  144 

zarhinus,  144 
variabilis 

Sciurus,  107 
varilla 

Suncus,  139 
uege"f!/.s 

Oligoryzonujs,  123 

Oryzomys,  123 
venustus 

Tamias,  109 

Thamnomys,  120 
verecundus 


Type  Specimens  of  Recent  Mammals  •  Helpen  and  McFadden 


181 


Stenoinys,  120 
Vespertilionidae,  145 
vexillariiis 

Paraxerus,  106 
vicinus 

Sciunis,  106 
victoriae 

Fitnisciiinis,  104 
vir 

Sorex,  139 
virgatus 

Agouti,  130 
virginiana 

Didelphis,  100 
virginianus 

Odocoileus,  148,  149 
vison 

Mnstela,  153,  154 

Putoriiis,  153 


Viverridae,  155 
volans 

Glaucomys,  104,  105 

Sciuroptenis,  104,  105 
vidcanitis 

Reithrodoutomi/s,  127 
vidgivaga 

Mustela,  154 
vtdgivagus 

Lutreola,  154 

Putorius,  154 
vidpes 

Vtdpes,  150,  151 
Vidpes 

bnngsi,  151 

deletrix,  150 

pennsylvanica,  150,  151 

ruhricatus,  151 

nibricosa,  150,  151 


ufl/ra,  150,  151 

vulpes,  150,  151 
vulpinus 

Sciun.is,  106 
ivallacei 

Myoictis,  100 
wavicus 

Myoictis,  100 
whijtei 

Cryptomys,  129 
woodfordi 

Pteropus,  141 
xanthogenys 

Putorius,  154 
xerampelinus 

Akodon,  127 

Scotinoniys,  127 
yucatanensis 

Sciurus,  106 


zappeyi 

Epirnys,  119 

Zapus 

acadicus,  112 
luirdyi,  112 
hudsonicus,  112 
ladas,  112 
orarius,  112 
trinotatus,  112 

zarhinus 

Vampyrops,  144 

zibethicus 
Fiber,  116 
Ondatra,  116 

Zygodontomys 
brevicauda,  128 
cheniei,  128 
seorsus,  128 


(US  ISSN  0027-4100) 


OF     THE 


seum 


A  New  Species  of  Lizard  Related  to 
Stenocercus  caducus  (Cope)  (Squamata: 
Iguanidae)  From  Peru  and  Bolivia,  With  a 

Key  to  the  "Ophryoessoides  Group" 


JOHN  E.  CADLE 

IJBRARY 

OCT  2  3  2002 

HARVARD 

HARVARD  UNIVERSITY 

CAMBRIDGE,  MASSACHUSEI  IS,  U.S.A. 

VOLUME  157,  NUMBERS 
14  November  2001 

(US  ISSN  0027-4100) 


PUBLICATIONS  ISSUED 

OR  DISTRIBUTED  BY  THE 

MUSEUM  OF  COMPARATIVE  ZOOLOGY 

HARVARD  UNIVERSITY 


Breviora  1952- 

bulletin  1863- 

Memoirs  1865-1938 

JoHNSONiA,  Department  of  Mollusks,  1941-1974 

Occasional  Papers  on  Mollusks,  1945- 

SPECIAL  PUBLICATIONS. 

1.  Whittington,  H.  B.,  and  W.  D.  I.  Rolfe  (eds.),  1963  Phylogeny  and 
Evolution  of  Crustacea.  192  pp. 

2.  Turner,  R.  D.,  1966.  A  Survey  and  illustrated  Catalogue  of  the  Tere- 
dinidea  (Mollusca:  Bivalvia).  265  pp. 

3.  Sprinkle,  J.,  1973.  Morphology  and  Evolution  of  Blastozoan  Echino- 
derms.  284  pp. 

4.  Eaton,  R.  J.,  1974.  A  Flora  of  Concord  from  Thoreau's  Time  to  the 
Present  Day.  236  pp. 

5.  Rhodin,  A.  G.  J.,  and  K.  Miyata  (eds.),  1983.  Advances  in  Herpetology 
and  Evolutionary  Biology:  Essays  in  Honor  of  Ernest  E.  Williams. 

725  pp. 

6.  Angelo,  R.,  1990.  Concord  Area  Trees  and  Shrubs.  118  pp. 

Other  Publications. 

Bigelow,  H.  B.,  and  W.  C.  Schroeder,  1953.  Fishes  of  the  Gulf  of  Maine. 
Reprinted  1964. 

Brues,  C.T.,  A.  L.  Melander,  and  F.  M.  Carpenter,  1954.  Classification  of 
Insects.  (Bulletin  of  the  M.  C.  Z,  Vol.  108.)  Reprinted  1971. 

Creighton,  W.  S.,  1950.  The  Ants  of  North  America.  Reprinted  1966. 

Lyman,  C.  P.,  and  A.  R.  Dawe  (eds.),  1960.  Proceedings  of  the  First  In- 
ternational Symposium  on  Natural  Mammalian  Hibernation.  (Bulletin 
of  the  M.  C.  Z,  VoL  124.) 

Orinthological  Gazetteers  of  the  Neotropics  (1975-). 

Peter's  Check-list  of  Birds  of  the  World,  vols.  1-16. 

Proceedings  of  the  New  England  Zoological  Club  1899-1947.  (Complete 
sets  only.) 

Proceedings  of  the  Boston  Society  of  Natural  History. 

Price  list  and  catalog  of  MCZ  publications  may  be  obtained  from  Publica- 
tions Office,  Museum  of  Comparative  Zoology,  Harvard  University,  Cambridge, 
Massachusetts  02138,  U.S.A. 

This  publication  has  been  printed  on  acid-free  pennanent  paper  stock. 

©The  President  and  Fellows  of  Harvard  College  2001. 


A  NEW  SPECIES  OF  LIZARD  RELATED  TO  STENOCERCUS 
CADUCUS  (COPE)  (SQUAMATA:  IGUANIDAE)  FROM  PERU  AND 
BOLIVIA,  WITH  A  KEY  TO  THE  "OPHRYOESSOIDES  GROUP " 


JOHN  E.  CADLE1 

CONTENTS 


Abstract  --  183 

Resumen 183 

Introduction  184 

Materials  and  Methods  184 

Description  of  a  New  Species  of  Stenocersus  187 

Stenocercus  prionotus  new  species  _  187 

Diagnosis  and  Comparisons   191 

Description  „ 193 

Distribution  Patterns  in  Stenocercus  prionotus 

197 

Patterns  of  Sympatry  and  Geographic  Variation 

in  Stenocercus  prionotus,  and  the  Need  for 

Additional  Fieldwork 200 

Natural  Histoiy  of  Stenocercus  prionotus  205 

Comparison  of  Stenocercus  prionotus  with  S. 

caducus  206 

Distributions  of  Stenocercus  prionotus  and  S. 

caducus  in  Eastern  Bolivia  210 

Is  the  Distribution  of  Stenocercus  aculeatus 

disjunct?  212 

Key  to  Species  of  the  " Ophryoessoides  group" 

of  Stenocercus  212 

Acknowledgments  215 

Appendix:  Specimens  Examined  216 

Literature  Cited  218 

Abstract.  A  new  species  of  iguanid  lizard,  Steno- 
cercus prionotus,  is  described  from  eastern  Peiii  and 
Bolivia  (known  range  froin  San  Martin  Department, 
Peru,  to  northern  La  Paz  and  El  Beni  departments, 
Bolivia).  Most  localities  are  in  the  Andean  foothills 
and  immediately  adjacent  lowlands.  Stenocercus 
prionotus  is  similar  to  several  other  species  of  Sten- 
ocercus with  large  posterior  head  scales,  an  enlarged 
row  of  supraoculars,  and  keeled  ventral  scales.  These 
similar  species  are  referred  to  as  the  "Ophryoessoides 
group"  without  implying  that  it  is  a  monophyletic  as- 
semblage. Based  on  their  common  possession  of  a 


'  Department  of  Herpetology,  Chicago  Zoological 
Society,  3300  Golf  Road,  Brookfield,  Illinois  60513. 
Research  Associate,  Department  of  Herpetology, 
Museum  of  Comparative  Zoology. 


unique  scaly  flap  concealing  a  portion  of  the  posthu- 
meral  mite  pocket,  the  new  species  is  apparently 
closely  related  to  S.  caducus  (Cope),  which  is  known 
from  central  Bolivia  south  to  northern  Argentina  and 
Paraguay.  Stenocercus  prionotus  is  distinguished  from 
S.  caducus  by  having  a  more  prominent  vertebral 
crest  and  a  pattern  of  alternating  light  and  dark  bars 
on  the  throat  (rarely  obseived  in  S.  caducus,  which 
usually  has  light  throat  spots).  These  two  species  also 
occupy  different  physiographic  regions  (western  Am- 
azonian rainforest  for  S.  prionotus;  chaco  for  S.  cad- 
ucus). 

Populations  of  S.  prionotus  from  northern  Peru 
have  a  higher  vertebral  crest  tlian  those  in  southern 
Peru  and  Bolivia.  Northern  populations  are  also 
broadly  sympatric  with  two  other  species  of  the 
"Ophryoessoides  group,"  S.  aculeatus  and  S.  fimbria- 
tus.  However,  in  southern  Peru  S.  prionotus  is  not 
known  to  be  sympatric  with  other  species  of  that 
group.  I  postulate  that  the  higher  vertebral  crest  in 
northern  populations  of  S.  prionotus  functions  as  a 
species  recognition  signal  in  the  multispecies  assem- 
blages. A  key  to  species  of  the  "Ophryoessoides 
group"  is  provided  and  distributions  of  the  species  in 
Peru  and  Bolivia  are  summarized. 

Resumen.  Se  describe  una  nueva  especie  de  lagar- 
tija  iguanida,  Stenocercus  prionotus,  del  Peru  Orien- 
tal y  de  Boli\aa.  Se  conoce  la  nueva  especie  desde  el 
departamento  de  San  Martin,  Peni,  hasta  el  norte  de 
los  departamentos  La  Paz  y  El  Beni  en  Bolivia.  La 
mayoria  de  las  localidades  se  encuentran  en  las  estri- 
baciones  andinas  y  adyacentes  tierras  bajas.  Stenocer- 
cus prionotus  es  similar  a  varias  otras  especies  de 
Stenocercus  con  grandes  escamas  sobre  el  posterior 
de  la  cabeza,  una  fila  amplia  de  supraoculares,  y  es- 
camas ventrales  quilladas.  Se  refiere  estas  especies 
como  el  "grupo  Ophryoessoides,"  sin  implicar  su 
monophyletismo.  Basada  en  su  posesion  de  un  lobulo 
escamoso  unico  que  oculta  una  porcion  del  bolsillo 
antehumeral,  se  considera  la  nueva  especie  cercana- 
mente  relacionada  a  Stenocercus  caducus  (Cope),  que 
se  conoce  desde  Bolivia  central  hasta  el  norte  de  la 


Bull.  Mus.  Comp.  ZooL,  157(3):  183-222,  November,  2001         183 


184         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


Argentina  y  del  Paraguay.  Stenocercus  prionotus  se 
distingue  de  S.  caduciis  al  tener  una  cresta  vertebral 
mas  prominente  y  un  patron  de  alternativas  barras 
claras  y  oscuras  sobre  la  garganta  (un  patron  rara- 
mente  observado  en  S.  caducus,  que  usualmente  ti- 
ene  manchas  claras  sobre  la  garganta). 

Las  poblaciones  de  Stenocercus  prionotus  del  norte 
del  Peni  tienen  una  cresta  vertebral  mas  alta  que  la 
cresta  en  poblaciones  del  sur  de  Peru  y  Bolivia.  Las 
poblaciones  del  norte  de  Peru  tambien  son  amplia- 
mente  simpatricas  con  dos  otras  especies  del  "gnipo 
Ophryoessoicles,"  S.  aculeatus  y  S.  finibriatus.  Sin  em- 
bargo, en  el  sur  del  Peni  y  Bolivia  no  se  conocen  lo- 
calidades  donde  se  encuentra  S.  prionotus  simpatrica 
con  otras  especies  del  giaipo.  Postulo  que  la  cresta  ver- 
tebral mas  alto  en  las  poblaciones  nortefias  de  S.  prion- 
otus functiona  como  un  serial  para  reconocimiento  de 
especies  en  comunidades  donde  hay  varias  especies 
simpatricas.  Se  provee  una  clave  para  las  especies  del 
"grupo  Ophnjoessoides"  y  se  resumen  las  distribu- 
ciones  de  las  especies  Peruanas  y  Bolivianas. 

INTRODUCTION 

Stenocercus  sensu  Frost  (1992),  includ- 
ing the  nominal  genera  Ophryoessoicles  and 
Proctotretus,  is  a  moderately  diverse  assem- 
blage of  South  American  iguanid  lizards 
(sensu  Macey  et  al.,  1997;  Schulte  et  al., 
1998)  with  about  50  species  currently  rec- 
ognized. Most  of  the  species  are  in  the  An- 
des and  adjacent  lowlands  of  Colombia,  Ec- 
uador, and  Pei"u,  but  a  few  are  primarily 
Amazonian  or  have  distributions  in  the 
physiographically  diverse  terrain  south  of 
the  Amazon  basin.  Although  new  species  of 
Stenocercus  continue  to  be  discovered  in 
the  field,  others  have  been  known  from  old 
collections  and  are  only  now  being  de- 
scribed (e.g.,  Cadle,  1991,  1998;  Avila-Pi- 
res,  1995).  In  this  category  is  a  new  species 
from  the  lowlands  of  eastern  Peru  and  Bo- 
livia that  has  been  referred  erroneously  to 
the  names  aculeatus  O'Shaughnessy  (1879) 
or  caducus  Cope  (1862)  in  previous  litera- 
ture, and  associated  with  the  genera  Lei- 
ocephalus  or  Ophnjoessoides  before  the 
current  understanding  of  these  names 
came  into  use  (see  Etheridge,  1966;  Frost, 
1992).  Rodriguez  and  Cadle  (1990)  left  the 
new  species  nameless  in  a  checklist  pend- 
ing resolution  of  its  status.  The  new  species 
is  apparently  closely  related  to  Stenocercus 
caducus  (Cope)  and  is  described  herein. 


MATERIALS  AND  METHODS 

Frost  (1992;  see  especially  footnote  5) 
and  Cadle  (1991)  discussed  reasons  for  re- 
ferring new  species  such  as  the  one  de- 
scribed here  to  Stenocercus  Dumeril  and 
Bibron  sensu  lato  (including  Ophnjoesso- 
ides Dumeril  and  Proctotretus  Dumeril 
and  Bibron).  Externally,  the  new  species  is 
similar  to  those  species  that  Fritts  (1974) 
placed  in  Ophnjoessoides,  that  is,  those 
species  with  keeled  ventral  scales,  large 
posterior  head  scales  (usually  including 
well-differentiated  interparietal,  parietals, 
postparietals,  and  occipitals),  and  one 
moderately  to  greatly  enlarged  supraocular 
row.  In  addition  to  the  new  species  de- 
scribed here,  species  included  in  the 
''Ophnjoessoides  group"  are  aculeatus 
O'Shaughnessy,  1879;  caducus  Cope, 
1862;  dunierilii  Steindachner,  1867;  ery- 
throgaster  Hallowell,  1856;  finibriatus  Avi- 
la-Pires,  1995;  huancahamhae  Cadle, 
1991;  iridescens  Giinther,  1859;  liniitaris 
Cadle,  1998;  scapidaris  Boulenger,  1901; 
tricristatus  Dumeril,  1851;  and  two  un- 
described  species  noted  later  in  this  paper 
under  Key  to  Species  of  the  "Ophryoesso- 
ides  group"  o/ Stenocercus.  I  use  the  term 
''Ophnjoessoides  group"  as  a  convenience 
to  refer  to  this  group  of  phenotypically 
similar  species  without  implication  as  to  its 
status  as  a  monophyletic  or  nonmonophy- 
letic  assemblage  within  Stenocercus.  m 

General  descriptive  protocols  follow  Ca- 
dle (1991),  who  defined  terminology  of  the 
scales,  neck  folds,  and  mite  pockets  used 
herein,  based  in  part  on  Frost  (1992).  Bi- 
lateral scale  counts  (e.g.,  subdigitals)  were 
made  only  on  one  side  (the  left,  unless  it 
was  damaged),  except  for  the  holotype,  for 
which  both  left  and  right  counts  were  re- 
corded (1,  r).  A  summary  of  selected  scu- 
tellational  and  qualitative  characters  for 
the  new  species  and  similar  species  from 
eastern  Peru  and  Bolivia  is  presented  in 
Table  1. 

All  measurements  are  in  millimeters. 
The  abbreviation  SVL  refers  to  the  head- 
body  length,  from  snout  to  vent.  The  con- 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle 


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186         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


Figure  1.  Distribution  of  species  of  Stenocercus  empfnasized  in  tfiis  paper  (western  South  America,  Ecuador  to  Paraguay  and 
northern  Argentina).  Open  symbols  for  S.  fimbhatus  are  literature  records  from  Avila-Pires  (1995);  locality  for  S.  caducus  in 
northern  Argentina  is  the  southernmost  locality  in  Argentina  reported  by  Cei  (1993).  Otherwise,  all  records  are  based  on  spec- 
imens examined.  Numbered  localities  are  documented  or  suspected  cases  of  sympatry  referred  to  in  the  text  and  noted  in  the 
Appendix:  1,  Pampa  Hermosa;  2,  Pampa  Seca,  Rio  Mixiollo;  3,  Tingo  Maria;  4,  Manu  National  Park.  Upper  left  quadrant  outlined 
with  dotted  line  is  the  area  shown  in  greater  detail  in  Figure  2. 


figurations  of  neck  folds  and  mite  pockets 
vaiy  considerably  among  species  of  Sten- 
ocercus and  are  useful  in  distinguishing 
species.  The  most  important  qualitative 
characteristics  of  these  features  used  here- 
in are  the  following,  which  are  discussed 
more  fully  by  Cadle  (1991): 

Neck  and  Body  Folds  and  Crests.  In 
contrast  to  many  species  of  Stenocercus, 
neck  folds  are  usually  absent  or  weakly  de- 


veloped in  the  "Opliryoessoides  group." 
When  present,  they  are  better  character- 
ized as  crests  rather  than  folds  because 
they  are  usually  indicated  by  strongly 
keeled  rows  of  scales  instead  of  actual 
folds  of  skin.  The  position  of  such  crests 
corresponds  to  the  position  of  folds  seen 
in  other  species  of  Stenocercus,  but  only 
two  are  commonly  seen  in  the  "Ophryoes- 
soides  group":  an  antehumeral  crest,  which 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle 


187 


is  a  more  or  less  vertical  row  of  strongly 
keeled  scales  immediately  anterior  to  the 
forelimb  insertion,  and  usually  highlighted 
with  white  scales;  and  a  supra- auricular 
crest,  a  strongly  keeled  horizontal  row  of 
scales  extending  from  the  posterior  tem- 
poral region  to  the  shoulder  region.  All 
species  of  the  ^'Ophnjoessoides  group" 
have  a  distinct  vertebral  crest  formed  by 
the  strongly  keeled,  and  often  projecting, 
scales  of  the  vertebral  row.  In  addition, 
some  of  these  species  have  a  more  or  less 
prominent  dorsolateral  crest  formed  by  a 
row  of  strongly  keeled  scales  separating 
the  flanks  from  the  dorsum  proper.  The 
dorsolateral  crest  varies  in  length.  In  some 
species  it  is  exceptionally  long,  extending 
from  the  proximal  portion  of  the  tail  to  the 
neck  region,  where  it  is  confluent  with  the 
supra-auricular  crest.  In  other  species  it  is 
present  only  anteriorly  or  posteriorly. 

Posthiimeral  (Axillary)  and  Postfemoral 
Mite  Pockets.  Type  1 — pocket  absent;  no 
skin  modification.  Type  2 — rudimentaiy 
pocket  manifested  by  skin  modification, 
such  as  bare  skin,  a  series  of  wrinkles,  or 
a  shallow  depression  lined  with  scales  dif- 
ferent from  surrounding  body  scales.  Type 
3 — similar  to  Type  2,  but  with  an  over- 
hanging fold  of  skin  or  a  thickened  border. 
Type  4 — a  deep  pocket,  usually  with  a 
broad  circular  opening,  whose  depth  is 
greater  than  half  the  diameter  of  its  open- 
ing. Type  5 — a  deep  pocket  with  a  narrow, 
slit-like  opening  and  a  depth  greater  than 
half  the  diameter  of  its  opening.  In  two 
species  discussed  herein  the  posthumeral 
pocket  is  partially  concealed  by  a  scaly  flap 
of  skin,  which  I  term  a  posthumeral  or  ax- 
illaiy  flap.  This  structure  is  described  more 
fully  later. 

Angulate  temporal  scales  are  distinctly 
enlarged,  keeled  scales  posterior  to,  and  in 
line  with,  the  superciliaiy  scales.  When 
present,  they  form  a  distinct  border  be- 
tween the  posterior  head  scales  and  the 
lateral  temporal  scales,  and  they  are  mor- 
phologically distinguishable  from  these  se- 
ries (Cadle,  1991:^6-7;  see  Fig.  4).  Angu- 
late temporal  scales  are  equivalent  to  su- 


pratemporals  as  used  in  some  literature 
(e.g.,  Avila-Pires,  1995).  In  several  species 
of  Stenocercus  the  angulate  temporals  are 
not  only  keeled  but  they  bear  a  projecting 
bladelike  vane  from  the  keels;  in  such  cas- 
es I  refer  to  the  scales  as  "projecting." 

Coordinates  for  localities  were  obtained 
from  the  ornithological  gazetteers  of  the 
Neotropics  (Stephens  and  Traylor,  1983; 
Paynter,  1989,  1992,  1993,  1997),  and 
from  Lamas  (1976),  Morales  and  Mc- 
Diarmid  (1996),  Schulenberg  and  Awbrey 
(1997b),  and  Peruvian  department  maps 
produced  by  the  Instituto  Geografico  Na- 
cional,  Lima.  I  also  consulted  the  on-line 
versions  of  the  Peru  and  Bolivia  gazetteers 
of  the  U.S.  Board  on  Geographic  Names 
at  the  GEOnet®  Names  Sei-ver:  http:// 
164.214.2.59/gns/html/index.html.  Brack- 
eted data  in  localities  are  inferences  from 
these  or  other  cited  sources.  Distributions 
of  the  new  species  and  others  emphasized 
in  this  paper  are  given  in  Figures  1  and  2. 
Institutional  abbreviations  are  given  at  the 
beginning  of  the  Appendix. 

DESCRIPTION  OF  A  NEW  SPECIES  OF 
STENOCERCUS 

Stenocercus  prionotus^ 
new  species 
Figures  3-7,  Figure  12;  Table  1 

Liocephalus  cadiicus  (Cope,  1862):  Boulenger  (1898), 
specimen  from  "Barraca,  Rio  Madidi"  [Boli\da] 
(BMNH  98.6.9.4)  and  probably  two  other  northern 
BoHvian  locahties  discussed  in  the  text. 

Ophnjoessoldes  caducus  (Cope,  1862):  Fugler  (1989: 
63),  specimens  from  San  Marcos  Ranch,  El  Beni 
Department,  Boli\da,  including  ROM  12815. 

Ophryoessoides  acideatus  (O'Shaughnessy,  1879): 
Fugler  (1983,  1986,  1989),  specimens  from  Tumi 
Chucua,  El  Beni  Department,  Bolix-ia  (USNM  par- 
atypes). 

Ophryoessoides  sp.:  Rodriguez  and  Cadle  (1990), 
specimen  from  Cocha  Cashu,  Manu  National  Park, 
Madre  de  Dios  Department,  Peru  (MCZ  150243). 


-  Stenocercus  pnonotus  was  recognized  as  new  by 
R.  Etheridge,  P.  E.  Vanzolini,  and  E.  E.  Williams 
many  years  ago.  Vanzolini  and  Williams  applied  the 
unpublished  name  Stenocercus  dorsatus  to  labels  of 
many  specimens  in  various  collections. 


188         Bulletin  Museum  of  Comparative  Zoologij,  Vol.  157,  No.  3 


7511^0° 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle        189 


Figure  3.     Dorsal  and  ventral  views  of  the  holotype  of  Stenocercus  prionotus  (USNM  193683).  Approximately  xO.87. 


Figure  2.  Northern  Peru  and  Ecuador  (see  Fig.  1)  showing  distributions  of  species  in  the  "Ophryoessoides  group."  Numbered 
localities  are  documented  or  suspected  cases  of  sympatry  referred  to  in  the  text  and  the  Appendix  (see  Fig.  1  for  names).  Open 
circle  at  06°S,  77°W  is  the  type  locality  for  Stenocercus  aculeatus  (Moyabamba,  San  Martin  Department).  All  other  localities  are 
based  on  specimens  examined;  see  Cadle  (1991,  1998)  for  S.  iridescens,  S.  limitaris,  and  S.  huancabambae.  The  known 
distributions  of  all  species  are  indicated  by  the  localities  plotted;  however,  the  distribution  of  S.  /r/descens  continues  farther  north 
in  western  Ecuador  than  the  area  covered  by  the  map. 


190 


Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


Holotype  (Figs.  3—5).  United  States 
National  Museum  of  Natural  History 
(USNM)  193683  (field  number  WCS 
2421).  PERU:  Depto.  Huanuco:  Jardin 
Botanico  de  la  Universidad  Agraria  de  la 
Selva,  Tingo  Maria,  vicinity  of  Rio  Hual- 
laga,  670  m  elevation  [09°18'S,  75°59'W]. 
Adult  male  collected  29  June  1966  by 
Wade  C.  Sherbrooke. 

Faratypes  from  the  Vicinity  of  the  Type 
Locality.  PERU:  Depto.  Huanuco:  Ca.  2 
mi.  by  trail  W.  Tingo  Maria,  west  bank  of 
Rio  Huallaga  in  vicinity  of  confluence  with 
Rio  Monzon,  670  m  elevation  (9  October 
1966,  W.  C.  Sherbrooke),  USNM  193685. 
Universidad  Agraria  de  la  Selva,  Tingo 
Maria,  Rio  Huallaga,  670  m  elevation  (18 
August  1967,  unknown  collector  for  W.  C. 
Sherbrooke),  USNM  193686.  Vicinity  of 
Cueva  de  las  Lechuzas,  ca.  3  mi.  SW  Tingo 
Maria,  Rio  Monzon,  ca.  700  m  elevation 
(17  April  1968,  Vito  Yaringano  for  W.  C. 
Sherbrooke),  USNM  193687.  Picuriacu, 
ca.  2  mi.  NW  Tingo  Maria,  Rio  Huallaga 
(20  April  1968,  W.  C.  Sherbrooke),  USNM 
193688. 

Other  Faratypes.  PERU:  Depto. 
Huanuco:  Buena  Vista,  Valley  of  the 
Chimchao  [  =  Rio  Chinchao]  [approximate- 
ly 9°31'S,  75°52'W]  (1-15  September 
1923,  E[dmund]  Heller),  FMNH  5582- 
83.  Hacienda  Pampayacu  [09°33'S, 
75°54'W]  (17  July-16  August  1936,  Dr. 
Snonge),  MCZ  43758-59,  43761-62. 
Rio  Llullapichis,  4—5  km  upstream  from 
Rio  Pachitea,  200  m  elevation  [09°37'S, 
74°55'W]  (Januaiy  1969,  Hans  W.  Koep- 
cke),  KU  179058.  [Depto.  Loreto]: 
E  [astern]  Peru,  Pampa  Hermosa,  near 
mouth  of  Rio  Cushabatay,  Rio  Ucayali  Val- 
ley 500  ft.  [152  m]  [07°12'S,  75°17'W] 
(date  unknown,  H[ai-vey]  Bassler),  AMNH 
56760-64.  Depto.  Madre  de  Dios:  Co- 
cha  Cashu  Biological  Station,  Manu  Na- 
tional Park  [11°51'S,  71°19'W]  (July  1975, 
John  W.  Fitzpatrick),  MCZ  150243.  Ex- 
plorer's Inn,  Tambopata  Reserve,  ca.  30 
km  (straight  line)  SSW  Puerto  Maldonado, 
280  m  [12°50'S,  69°17'W]  (2  September- 
7  October  1983,  native  collectors),  USNM 


247468-69,  247680;  (23  May  1986,  Vic- 
tor R.  Morales),  USNM  269022.  Depto. 
Puno:  Prov.  Sandia:  Tambopata,  San  Juan 
[del  Oro],  1520  m  or  5000  ft.  [14°12'S, 
69°08'W]^  (22  November-20  December 
1950,  Hilda  H.  Heller),  FMNH  64788- 
92,  64794-811.  [Depto.  San  Martin]: 
Juanjui  [07°11'S,  76°45'W]  (collector  and 
date' unknown),  MCZ  121233.  Tarapoto, 
370  m  [06°30'S,  76°25'W]  (25  July  1984, 
Rainer  Schulte),  KU  212629. 

ROLIVIA:  Depto.  Reni:  Provincia  Va- 
cadiez,  Tumi  Chucua  [176  m;  11°08'S, 
66°10'W]  (23  October-18  November 
1981,  Charles  M.  Fugler),  USNM 
280246—51.  Puerto  Cruzeiro,  San  Marcos 
Ranch  at  confluence  of  Rios  Isiboro  and 
Ichoa  [15°17'S,  65°45'W]  (10  February 
1977,  J.  Lovisek),  ROM  12815.  [Depto. 
La  Paz]:  Barraca,  Rio  Madidi  [12°35'S, 
67°02'W]  (1893,  Luigi  Balzan),  RMNH 
98.6.9.4. 

Distribution  (Fig.  1).  Stenocercus  prion- 
otiis  is  known  from  the  lowlands  and  An- 
dean foothills  of  eastern  Peru  and  adjacent 
Bolivia  (San  Martin  department,  Peru, 
south  to  the  Rio  Beni  in  northern  Bolivia). 
The  known  elevational  range  is  176—1,520 
m,  but  the  highest  recorded  elevation  is 
more  than  twice  the  elevation  of  the  next 
lower  one.  Most  localities  in  Peru  are  ad- 
jacent to  or  near  the  Andean  foothills,  and 
several  localities  in  La  Paz  and  El  Beni 
departments,  Bolivia,  are  unconfirmed.  No 
specimens  are  known  from  a  broad  geo- 
graphic hiatus  between  central  and  south- 
ern Peru  (Fig.  1).  See  Distribution  Fat- 
terns  in  Stenocercus  prionotus  for  further 
discussion. 

Etymology.  The  epithet  prionotus  is  de- 


^  Hilda  Hellers  notes  on  this  collection  in  the  Karl 
P.  Schmidt  archives  of  the  FMNH  describe  San  Juan 
as  the  site  of  an  agricultural  station  on  "the  left  side 
of  die  Rio  Tambopata  at  5000  feet."  With  some  hes- 
itancy I  identify  Heller's  locality  as  the  town  kno\\'n 
as  San  Juan  del  Oro,  which  is  on  the  left  bank  of  the 
Rio  Tambopata  at  approximately  the  elevation  given 
by  Heller.  I  have  located  only  two  other  places  named 
San  Juan  in  Puno  Department,  but  neither  is  on  the 
Rio  Tambopata. 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle 


191 


rived  from  the  Greek  adjective  prionotos 
meaning  jagged  or  serrate.  The  name  re- 
fers to  the  strongly  serrate  vertebral  crest 
of  Stenocercus  prionotus,  which  is  the 
most  obvious  character  distinguishing  this 
species  from  its  apparent  closest  relative, 
S.  caduciis. 

Data  on  the  Holotype.  Adult  male,  hem- 
ipenes  partially  everted.  SVL,  83  mm.  Tail 
length,  201  mm.  Total  length,  284  mm. 
Tail/total  length,  0.71.  Vertebral  scales  be- 
tween the  occipital  and  the  posterior  mar- 
gin of  the  hind  limb,  31.  Midbody  scales, 
42.  Gular  scales  between  the  ears,  16.  In- 
ternasals,  6.  Subdigital  scales  on  fourth  fin- 
gers and  toes,  respectively,  18—18,  25—25. 
Color  pattern  well  presei-ved:  top  of  head 
brown  with  narrow  dark  brown  interorbital 
bar  extending  laterally  onto  supraoculars; 
dorsum  brown  with  narrow  blackish  chev- 
rons middorsally  (1  on  neck,  1  above  fore- 
limbs,  2  others  anterior  to  midbody;  pos- 
terior chevrons  poorly  defined);  dark 
brown  scapular  blotch  bordered  anteriorly 
by  white  antehumeral  stripe;  ill-defined 
grayish  dorsolateral  streaks  between  ear 
and  anterior  body;  throat  grayish  with 
poorly  defined  oblique  light  grayish 
stripes;  venter  brown  without  distinct  pat- 
tern. 

Definition.  A  species  of  Stenocercus 
characterized  by  the  following  features:  (1) 
Dorsal  head  scales  subimbricate  and 
strongly  keeled  to  multicarinate;  temporals 
keeled,  imbricate  or  subimbricate.  (2)  Pos- 
terior head  scales  larger  than  anterior 
ones,  with  distinct  interparietal,  a  pair  of 
parietals,  a  pair  of  postparietals,  and  a 
large  median  occipital  (often  surrounded 
by  several  small  irregular  scales).  (3)  In- 
ternasals  usually  7,  but  pattern  irregular 
and  may  be  5  or  6.  (4)  One  row  of  supra- 
oculars distinctly  enlarged.  (5)  One  canthal 
on  each  side  between  the  superciliaries 
and  the  lateralmost  internasal.  (6)  A  pair 
of  strongly  keeled  angulate  temporals  in 
line  on  each  side  (rarely,  3  angulate  tem- 
porals are  present),  each  with  a  low  pro- 
jecting blade;  partially  or  completely  sep- 
arated from  enlarged  posterior  head  scales 


by  a  single  row  of  small  scales.  (7)  Anterior 
and  posterior  gular  scales  strongly  keeled. 
(8)  Parietal  eye  distinct.  (9)  Neck  folds  ab- 
sent; a  vertical,  strongly  keeled  row  of 
scales  in  the  antehumeral  region  and  oc- 
casionally a  much  less  distinct  raised  series 
in  the  supra-auricular  region.  (10)  Dorsal 
and  ventral  body  scales  imbricate,  mucro- 
nate,  strongly  keeled;  dorsal  scales  at  mid- 
body  36—48.  (11)  Vertebral  row  continu- 
ous, bearing  a  strongly  projecting  serrate 
crest  in  adults;  a  dorsolateral  crest  present 
on  posterior  body  and  the  base  of  the  tail. 

(12)  Deep  posthumeral  pocket  (Type  4) 
partially  concealed  by  a  scaly  posthumeral 
flap  originating  on  its  anteroventral  bor- 
der; postfemoral  pocket  absent  (Type  1). 

(13)  Scales  of  posterior  thigh  imbricate, 
keeled.  (14)  Tail  strongly  compressed  in 
adults,  anteriorly  with  low  vertebral  and 
dorsolateral  crests  continuous  with  those 
of  the  body.  (15)  Dorsal  coloration  of 
males  in  preservative  (Figs.  3,  6)  brown 
with  or  without  distinct  chevrons;  a  dis- 
tinct white  vertical  antehumeral  stripe  ex- 
tending ventrally  to  the  proximal  ventral 
surface  of  forelimb;  a  large  dark  scapular 
blotch;  in  well-presei"ved  specimens  the 
throat  bears  oblique  alternating  dark  and 
light  stripes  (see  Description);  females 
similar  but  pattern  elements  often  more 
subdued. 

DIAGNOSIS  AND  COMPARISONS 

In  having  enlarged  posterior  head 
scales,  an  enlarged  row  of  supraoculars, 
and  strongly  keeled  ventral  scales,  Steno- 
cercus prionotus  is  like  other  species  in  the 
"Ophryoessoides  group"  of  Stenocercus. 
These  are  the  species  most  likely  to  be 
confused  with  S.  prionotus.  Five  other  de- 
scribed species  of  the  ''Ophryoessoides 
group"  occur  in  eastern  Peru  or  Bolivia:  S. 
aculeatus  (O'Shaughnessey,  1879);  S.  cad- 
ucus  (Cope,  1862);  S.  finibriatus  Avila-Pi- 
res,  1995;  S.  huancabanibae  Cadle,  1991; 
and  S.  scapularis  (Boulenger,  1901).  An 
undescribed  species  occurs  in  the  Rio 
Maraiion  valley  of  eastern  Peru  (see  Key 
to  Species  of  the  "Ophryoessoides  Group" 


192         Bulletin  Museum  of  Comparative  Zoologtj,  Vol.  157,  No.  3 


of  Stenocercus).  Stenocercus  prionotus 
and  S.  caducus  are  unique  among  known 
species  of  Stenocercus  (perhaps  unique 
within  iguanids)  in  having  deep  posthu- 
meral  mite  pockets  (Type  4)  that  are  par- 
tially concealed  anteroventrally  by  a  scaly 
flap,  which  may  be  termed  a  posthumeral 
or  axillary  flap  (Fig.  5).  Stenocercus  prion- 
otus and  S.  caducus  are  compared  in 
greater  detail  below,  but  S.  pmonotus  is 
distinguished  from  S.  caducus  (character- 
istics in  parentheses)  by:  (1)  a  strongly  pro- 
jecting, serrate  vertebral  crest  (low  and 
scarcely  projecting);  (2)  2  (usually)  or  3  en- 
larged, strongly  keeled,  projecting  angu- 
late  temporal  scales  on  each  side  (scales 
not  greatly  enlarged,  less  projecting);  and 
(3)  a  gular  pattern  consisting,  when  evi- 
dent, of  oblique  alternating  dark  and  light 
lines  or  bars,  or  oblique  light  lines  on  a 
dark  ground  color  (usually  light  spots  on  a 
darker  ground  color,  unicolor,  or  [rarely]  a 
pattern  similar  to  that  of  S.  prionotus). 

Readily  determined  characters  distin- 
guishing Stenocercus  prionotus  from  the 
other  four  species  of  the  "Ophryoessoides 
group"  known  from  eastern  PeiTi  and  Bo- 
livia include  the  extent  of  keeling  on  dorsal 
head  and  body  scales,  relative  develop- 
ment of  the  postfemoral  pockets,  and  the 
number  of  midbody  scale  rows  (Table  1 
and  key  presented  later  herein).  Stenocer- 
cus fimbriatus  and  S.  aculeatus  are  known 
to  be  sympatric  with  S.  prionotus  at  several 
localities  in  eastern  Peru.  In  addition  to 
having  a  posthumeral  flap  (absent  in  S. 
fimbriatus  and  S.  acideatus),  S.  prionotus 
is  distinguished  from  S.  fimbriatus  (char- 
acteristics in  parentheses;  see  Avila-Pires, 
1995)  in  having  strongly  keeled  dorsal 
scales  in  adults  (smooth  or  weakly  keeled), 
a  dorsolateral  crest  prominent  only  on  the 
posterior  body  (prominent  anteriorly  and 
continuous  with  antehumeral  and  supra- 
auricular  folds  or  crests),  and  in  lacking 
"fimbriate"  scales  on  the  posterior  distal 
portion  of  the  thigh  (present).  Stenocercus 
prionotus  is  distinguished  from  S.  aculea- 
tus (characteristics  in  parendieses)  in  lack- 
ing   a   postfemoral    pocket    (moderate   to 


deep);  in  having  strongly  keeled,  often 
multicarinate,  head  scales  (smooth  or 
weakly  striated  in  adults,  wrinkled  in  ju- 
veniles); 5—7  internasals  (4—5);  and  only 
moderately  enlarged  supraoculars,  usually 
5—6  supraoculars  across  the  widest  part  of 
the  orbit  (greatly  enlarged,  usually  4  across 
the  orbit). 

Stenocercus  prionotus  differs  from  S. 
scapularis  (characteristics  in  parentheses) 
in  lacking  squarish  or  rectangular  project- 
ing superciliaiy  scales  (present)  and  in 
having  fewer  than  50  midbody  dorsal  scale 
rows  (59—70  rows).  Stenocercus  prionotus 
differs  from  S.  huancabambae  (characters 
in  parentheses)  in  lacking  a  postfemoral 
pocket  (deep,  Type  5)  and  in  having  prom- 
inent dorsolateral  crests  on  the  posterior 
body  (weak,  restricted  to  anterior  body 
when  present). 

Two  species  of  Stenocercus  from  west- 
ern Ecuador  and  Peru,  S.  iridescens  and 
S.  limitaris,  have  enlarged  posterior  head 
scales  and  supraoculars.  In  contrast  to 
Stenocercus  prionotus,  S.  iridescens  has 
smooth  head  plates,  2  canthals,  a  poorly 
developed  posthumeral  pocket  (Type  1  or 
2),  and  lacks  keeled  angulate  temporals 
and  dorsolateral  crests  on  the  body  (see 
Cadle,  1991,  fig.  10).  Stenocercus  limitaris 
has  a  deep  postfemoral  pocket  (Type  5),  2 
canthals,  a  single  strongly  keeled  (but  non- 
projecting)  angulate  temporal,  and  lacks 
dorsolateral  crests. 

Other  non-Peruvian  species  of  the 
"Ophryoe.ssoides  group"  can  be  distin- 
guished from  Stenocercus  prionotus  by 
features  in  the  key  presented  later  and 
other  superficial  characters,  such  as  the 
presence  of  2  canthals  and  4  internasals 
(usually)  in  S.  erythrogaster  (1  and  5—7  in 
S.  prionotus),  and  enlarged,  projecting  py- 
ramidal or  conical  postsuperciliary  scales 
in  S.  dumerilii  and  S.  tricristatus  (Avila- 
Pires,  1995). 

Other  species  of  Stenocercus  are  distin- 
guished from  S.  prionotus  by  a  combina- 
tion of  features  such  as  smaller  head 
plates,  smooth  ventrals,  and  absence  of 
dorsolateral  crests.  Most  species  of  Steno- 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle 


193 


cercus  except  the  "Ophnjoessoides  group" 
have  smooth  or  weakly  keeled  head  plates. 
Other  characters,  such  as  the  number  of 
dorsal  scale  rows,  morphology  of  the  pos- 
thumeral  and  postfemoral  mite  pockets, 
extent  of  sexual  dimoi*phisin,  and  degree 
of  differentiation  of  the  vertebral  scale  row 
and  crest  also  aid  in  distinguishing  the  spe- 
cies (see  descriptions  and  discussion  in 
Fritts,  1974;  Cadle,  1991,  1998). 

Apart  from  Stenocercus  fimbriatus,  S. 
scapularis,  and  S.  acideatus,  two  other  spe- 
cies of  Stenocercus  are  known  from  local- 
ities close  to  or  synipatric  with  known  pop- 
ulations of  S.  prionotus:  S.  crassicaudatus 
and  S.  roseiventris.  These  species  are  dis- 
tinguished from  S.  prionotus  by  lacking 
prominent  serrate  vertebral  crests  (low 
crests  may  be  present),  having  smooth 
ventral  scales,  and  having  prominently  spi- 
nose  tails  with  the  spines  arranged  in  dis- 
tinct whorls. 

DESCRIPTION 

Head  (Fig.  4).  Dorsal  head  scales  sub- 
imbricate  (a  tendency  to  be  more  juxta- 
posed posteriorly);  strongly  keeled  to  mul- 
ticarinate  or  wrinkled.  Rostral  in  contact 
with  first  supralabial,  first  lorilabials,  and  a 
series  of  postrostrals.  Usually  7  elongate, 
strongly  keeled  internasals  between  the 
nasals  dorsally;  however,  the  anterior  dor- 
sal head  scales  are  very  irregular  and  oc- 
casionally only  5  or  6  internasals  are  pre- 
sent. One  canthal  scale  between  the  an- 
terior superciliary  and  the  lateralmost  in- 
ternasal,  separated  from  the  nasals  by  tiny 
postnasals.  Canthus  very  strongly  angled. 
Nostril  in  posterior  portion  of  an  elongate 
nasal  scale,  which  may  contact  the  rostral 
scale  anteriorly  or  be  separated  from  it  by 
small  postrostrals.  Four  or  5  strongly  over- 
lapping, elongate  anterior  superciliaries 
followed  by  2  or  3  shorter  posterior  su- 
perciliaries slightly  overlapping  in  the  re- 
verse direction  (but  more  or  less  in  a 
straight  line).  One  supraocular  row  inod- 
erately  enlarged,  2  mediocentral  scales 
much  larger  than  the  others.  Five  or  6 
scales   across   the   supraocular  area  at  its 


widest  part.  Interparietal  distinct  and  elon- 
gate, diamond-shaped  or  pentagonal  (apex 
posteriorly).  Parietal  eye  visible.  A  pair  of 
parietals  in  contact  behind  the  inteqDari- 
etal,  flanked  posterolaterally  by  a  postpar- 
ietal  on  each  side.  Postparietals  separated 
medially  by  a  single  median  transversely 
elongate  occipital;  occasionally  1  or  2  small 
scales  are  intercalated  at  the  juncture  of 
the  parietal,  postparietal,  and/or  occipital 
(e.g..  Fig.  4). 

Lateral  temporal  scales  strongly  keeled, 
imbricate  to  subimbricate;  separated  from 
posterior  dorsal  head  scales  on  each  side 
by  2  (occasionally  3)  elongate,  strongly 
keeled  angulate  temporal  scales  bearing  a 
low  projecting  vane.  Keels  of  adjacent  an- 
gulate teinporals  aligned.  Posterior  angu- 
late temporals  separated  from  postparie- 
tals by  2  or  3  small  scales  in  a  longitudinal 
row.  Anterior  angulate  temporals  may  con- 
tact postparietal  and  one  other  larger  pos- 
terior head  scale  or  be  separated  from 
them  by  small  scales.  Anterior  border  of 
ear  weakly  denticulated;  posterior  border 
rounded,  bordered  with  keeled  imbricate 
scales. 

Anterior  and  posterior  gulars  strongly 
keeled.  Mental  smooth,  in  contact  with 
first  pair  of  postmentals  and  first  pair  of 
infralabials.  Enlarged  postmentals  3  or  4 
on  each  side,  only  the  first  pair  in  contact 
medially. 

Neck  and  Body.  Dorsal  and  lateral 
scales  of  neck  and  body  imbricate,  mucro- 
nate,  strongly  keeled.  Vertebral  row  pro- 
duced into  a  prominent  projecting  serrate 
crest  in  adults  of  both  sexes  that  is  contin- 
uous from  the  nuchal  region  to  the  base 
of  the  tail,  gradually  disappearing  on  the 
anterior  Va  to  V2  of  tail.  Dorsolateral  crest 
(a  raised,  strongly  keeled  row  of  scales)  on 
posterior  %  of  body,  continuing  onto  base 
of  tail.  The  dorsolateral  crest  occasionally 
appears  very  indistinctly  farther  anteriorly 
on  the  body,  but  only  on  the  posterior 
body  does  it  shaiply  delimit  the  dorsolat- 
eral (paradorsal)  scales  from  the  flank 
scales.  Three  rows  of  scales  between  dor- 
solateral and  vertebral  crests  at  anterior 


194         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


Figure  4.  Head  scales  of  the  holotype  of  Stenocercus  prionotus  (USNM  193683)  in  dorsal  and  lateral  views.  Scale  bars  =  3 
mm.  To  facilitate  coordination  with  the  text,  the  following  scales  are  indicated  on  one  side  (interparietal  and  occipital  are  median 
scales);  A,  angulate  temporal;  I,  interparietal;  O,  occipital;  P,  parietal;  PP,  postparietal. 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Caclle 


195 


edge  of  dorsolateral  crest  (i.e.,  just  anterior 
to  the  pelvic  region),  2  rows  posteriorly 
(dorsal  to  hindlimbs);  scales  between  the 
crests  strongly  imbricate  and  keeled  only 
on  posterior  part  of  scale.  Flank  scales 
mostly  fully  keeled  (sometimes  only  the 
posterior  part  of  each  scale),  imbricate, 
mucronate,  slightly  smaller  than  dorsolat- 
eral scales.  Ventral  body  scales  mucronate, 
strongly  keeled  (keels  running  the  length 
of  the  scales).  Ventrals  approximately  the 
same  size  as  the  dorsolateral  scales,  larger 
than  flank  scales. 

Neck  Folds.  Distinct  neck  folds  absent. 
Poorly  developed  antehumeral  crest  pre- 
sent. 

Tail.  Tail  strongly  compressed,  anteriorly 
bearing  low  projecting  vertebral  and  dor- 
solateral crests  continuous  with  those  of 
the  body.  Dorsal  scales  moderately  keeled, 
ventral  scales  strongly  keeled. 

Limbs.  Dorsal  and  ventral  scales  of  fore- 
limbs,  hindlimbs,  and  posterior  thigh 
strongly  keeled,  unicarinate,  mucronate; 
some  scales  of  the  shank  larger  than  any 
thigh  scales.  Supradigitals  and  subdigitals 
unicarinate.  Palmar  scales  strongly  multi- 
carinate.  Plantar  scales  strongly  unicari- 
nate. 

Posfhiimeral  and  Postfemoral  Mite 
Pockets.  Posthumeral  mite  pocket  a  deep 
cavity  (Type  4)  with  a  prominent  axillaiy 
flap  concealing  the  anteroventral  aspect 
(Fig.  5).  Postfemoral  pocket  absent  (Type 

The  flap  associated  with  the  posthumer- 
al pocket  projects  from  the  anteroventral 
and  ventral  edges  of  the  pocket.  Antero- 
dorsally,  a  similar  but  much  smaller  flap  is 
present  in  some  specimens  (e.g..  Fig.  5). 
The  posthumeral  flap  consists  of  a  fleshy 
ridge  covered  anteriorly  and  posteriorly 
(or  externally  and  internally  when  the  flap 
is  lying  flat  against  the  body)  by  keeled  im- 
bricate scales.  Externally,  usually  3  or  4 
larger  scales  cover  the  flap  ventrally  and  a 
series  of  much  smaller  scales  is  present 
dorsally.  One  or  2  of  the  larger  scales  are 
sometimes  highlighted  with  white.  When 
appressed  against  the  body  (i.e.,   against 


Figure  5.  Posthumeral  (axillary)  flap  of  Stenocercus  priono- 
tus  (USNM  193683,  holotype).  Anterior  to  the  right.  The  broad 
oval  on  the  right  is  the  deflected  forelimb  and  the  posthumeral 
mite  pocket  is  the  heavily  stippled  cavity  deep  to  the  flap.  The 
posterior  border  of  the  axillary  flap  is  marked  by  the  scales 
with  heavily  outlined  posterior  borders  and  it  extends  anteriorly 
to  the  ventral  part  of  the  forelimb.  A  smaller  dorsal  flap  is  also 
present  in  this  specimen  (small  patch  of  heavily  outlined  scales 
on  anterodorsal  edge  of  the  pocket;  see  text).  Scale  bar  =  1 
mm. 


the  opening  of  the  posthumeral  pocket) 
the  flap  conceals  approximately  the  ventral 
Vs  to  Vi  of  the  vertical  dimension  of  the 
pocket.  The  flap  is  equally  prominent  in 
adults  of  both  sexes  and  is  proportionally 
as  well  developed  in  subadults  (including 
hatchlings)  as  in  adults. 

Size  and  Propoi~tions.  Largest  inale 
(USNM  193683)  89  mm  SVL,  323  mm  to- 
tal length  (sample  size  of  males  with  SVL 
>70  mm  =  13).  Largest  female  (KU 
212629)  93  mm  SVL,  329  mm  total  length 
(sample  size  of  females  with  SVL  >70  mm 
=  16).  Tail  relatively  long,  69-74%  of  total 
length  in  adults  (67-71%  in  juveniles). 

Coloration  and  Pattern  of  Adult  Males 
in  Life.  The  following  color  descriptions 
are  paraphrased  from  the  field  notes  of 
Wade  C.  Sherbrooke.  USNM  193683  (ho- 
lotype): 

Lower  half  of  side  between  limbs  is  lavender- 
brown.  This  color  extends  from  both  sides  across 
the  belly  approximately  Vs  of  the  way  on  each  side, 
leaving  a  tan-brown  central  strip  down  the  belly. 
General  base  color  of  the  body  is  brown,  darkest 


196         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


Figure  6.  Lateral  view  of  Stenocercus  prionotus  from  northern  Peru  (USNM  193685;  male,  snout-vent  length  76  mm).  Note 
the  high  vertebral  crest  and  the  following  pattern  elements:  dark  subocular  bar,  dark  blotches  on  the  dorsal  and  lateral  surfaces 
of  the  neck,  pale  antehumeral  bar,  indistinct  pale  dorsolateral  stripe,  and  indistinct  oblique  bars  on  the  trunk. 


near  the  hind  legs  and  tan  just  in  from  of  the  fore- 
legs. A  distinct  white  line  runs  dorsally  from  the 
top  of  each  foreleg  to  three  quarters  of  the  way  to 
the  dorsal  crest;  it  runs  through  a  large  black  patch 
just  above  the  forelimb.  Head  markings  consist  of 
a  dark  brown  line  i-unning  between  the  eyes  on  die 
top  of  die  head;  this  continues  through  the  eye  to 
broaden  slightly  at  the  rear  portion  of  the  jaw.  The 
gular  area  is  streaked  by  several  light  cream  lines. 
Very  slight  lavender  tinge  to  all  of  body  behind 
forelimbs.  [Sketch  in  notes  shows  black  middorsal 
patches  that  don't  extend  to  flanks]. 

USNM  193685  (WCS  2543): 

This  specimen  closely  resembles  [USNM  193683] 
in  color,  with  one  exception.  There  are  several 
green  spots  on  .  .  .  the  right  dorsal  surface  between 
the  limbs  and  on  the  dorsal  portions  of  the  tail 
behind  the  hind  limbs  and  the  dorsal  tibio-fibula 
portion  of  the  right  hind  leg. 

Coloration  of  Adult  Females  in  Life.  Un- 
known. 

Coloration  in  Preservative  (Figs.  3,  6,  7). 
Specimens  of  Stenocercus  prionotus  vary 
greatly  in  coloration  due  mainly  to  varia- 
tion in  initial  preservation  and  the  length 
of  time  in  presei-vative.  Poorly  preseived 
specimens  may  be  more  or  less  uniform 
brown  all  over,  although  obscure  pattern 
eleiTients  are  usually  present.  Well  pre- 
served specimens  are  brown  dorsally  with 
darker  brown  to  blackish  chevrons  mid- 
dorsally.  One  chevron  dorsal  to  each  pair 
of  limbs  and  one  on  the  neck  are  usually 
evident,  and  these  are  darker  than  others 
that  iTiay  be  present.  Three  to  five  mid- 
dorsal   chevrons   are   between   the   limbs. 


Dark  spots  or  an  additional  chevron  are 
often  present  on  the  dorsal  neck  and  usu- 
ally on  the  base  of  the  tail.  The  light  an- 
tehumeral/humeral  line  is  universally  pre- 
sent and  evidence  of  the  dark  shoulder 
patch  is  usually  present  (often  veiy  prom- 
inent). Flanks  usually  unicolor  and  some- 
what darker  than  the  dorsum  between  the 
dorsolateral  crests;  however,  soixie  speci- 
mens (e.g.,  KU  179058,  USNM  280246) 
have  distinct  dirty  white  vertical  bars  or 
chevrons  on  the  flanks  (five  between  the 
limbs),  and  such  bars  appear  occasionally, 
but  more  obscurely  in  other  specimens 
(see  Fig.  6).  The  dorsolateral  crest  is  often 
highlighted  for  a  variable  length  with  a  dis- 
tinct or  indistinct  light  line,  giving  the  im- 
pression of  a  light  dorsolateral  stripe. 
Forelimbs  more  or  less  unicolor  brown  or 
with  obscure  pattern;  hindlimbs  brown 
with  darker  brown  bands.  Dark  subocular 
bar  distinct.  Top  of  head  often  with  an  ob- 
scure or  distinct  dark  brown  interocular 
bar.  Oblique  bars  on  throat  (Fig.  7)  often 
visible  but  throat  may  be  unicolor  or  have 
an  obscure  pattern.  Venter  of  most  speci- 
mens unicolor,  dirty  white,  gray,  or  beige; 
however,  some  specimens  (e.g.,  USNM 
280246)  have  a  series  of  irregular  longi- 
tudinal dark  brown  streaks. 

Scale  Counts  and  Qualitative  Features 
(Table  1).  Stenocercus  prionotus  has  rela- 
tively low  midbody,  vertebral,  and  gular 
scale  counts.  The  scales  are  relatively  large 
and  strongly  keeled  over  most  of  the  body. 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle 


197 


Figure  7.  Gular  patterns  in  Stenocercus  prionotus.  Top,  typ- 
ical throat  pattern  (oblique  view)  consisting  of  light  and  dark 
stripes  that  extend  medially  to  the  midline  (USNM  193687). 
Bottom,  specimen  in  which  the  ground  color  is  lighter  and 
therefore  the  contrasting  pale  stripes  are  less  distinct  (USNM 
193685). 


Sexual  Dimorphism.  Stenocercus  prion- 
otus does  not  exhibit  strong  sexual  dimor- 
phism. Males  and  females  attain  approxi- 
mately the  same  size  and  have  the  same 
general  pattern,  but  whether  the  colora- 
tion in  life  reported  above  for  adult  males 
pertains  to  females  as  well  is  unknown. 
The  vertebral  crest  is  only  slightly  more 
developed  in  males  than  in  females  of  the 
same  population  but  this  character  shows 
strong  clinal  variation  (northern  popula- 
tions with  higher  crests;  further  discussed 
below).  Other  characters  that  sometimes 
vary  between  the  sexes  in  Stenocercus 
show  little  variation  in  S.  priontotus.  Nei- 


ther standard  meristic  counts  (Table  1)  nor 
the  relative  development  of  the  posthu- 
meral  and  postfemoral  pockets  (Types  4 
and  1,  respectively,  in  adults  of  both  sexes 
and  in  subadults)  show  obvious  sexual  di- 
morphism. 

DISTRIBUTION  PATTERNS  IN 
STENOCERCUS  PRIONOTUS 

Absence  of  Stenocercus  prionotus  from 
Lowland  Localities  in  Eastern  Fern.  Sten- 
ocercus prionotus  is  widespread  in  the 
lowlands  along  the  Andean  front  from 
northern  Peru  to  northern  Bolivia  (Fig.  1). 
However,  all  Peruvian  localities  are  close 
to  the  Andean  foothills  and  south  of  the 
broad  extension  of  the  Cordillera  Oriental 
separating  the  great  bend  of  the  Rio  Mar- 
aflon  from  upper  reaches  of  the  Rio  Hual- 
laga  (Fig.  2).  The  absence  of  specimens  in 
comprehensive  collections  from  the  Iqui- 
tos  region  (Dixon  and  Soini,  1986),  Balta 
(Ucayali  Department;  specimens  at 
LSUMNS  and  University  of  Arizona),  and 
Cuzco  Amazonico  (Madre  de  Dios  De- 
partment; Duellman  and  Salas,  1991)  sug- 
gest that  S.  prionotus  may  be  absent  from 
the  lowlands  distant  from  the  Andean  foot- 
hills, at  least  in  Peru.  Similarly,  collections 
from  northern  Loreto  Department  (Duell- 
man and  Mendelson,  1995)  and  northern 
Amazonas  Department  (J.  E.  Cadle  and  R. 
W.  McDiarmid,  unpublished  data  from  the 
Rio  Cenepa  and  Rio  Santiago)  suggest  that 
S.  prionotus  does  not  occur  north  of  the 
Rio  Marafion. 

However,  these  sites  have  been  sampled 
unevenly.  For  example,  species  accumula- 
tion curves  for  lizards  at  Cuzco  Amazonico 
(Duellman  and  Koechlin,  1991)  reached 
an  asymptote  after  about  15  person-weeks 
of  effort,  whereas  only  5  person-weeks 
were  expended  in  northern  Loreto  and  the 
species  accumulation  curve  for  the  total 
herpetofauna  showed  no  asymptote 
(Duellman  and  Mendelson,  1995;  data  not 
presented  for  lizards  only).  Quantitative 
data  are  not  available  for  the  other  sites, 
but  large  collections  are  available  for  the 
Iquitos  region,  including  more  than  1,000 


198         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


Table  2.  Patterns  of  presumed  sympatry  of  Stenocercus  in  eastern  Peru.  Only  species  of  the 
"Ophryoessoides  group"  and  the  superficially  similar  species,  S.  roseiventris,  are  listed. 1  Locali- 
ties ARE  listed  roughly  NORTH  TO  SOUTH;  FOR  PRECISE  LOCALITIES  SEE  THE  APPENDIX  AND  LOCALITIES 

FOR  THE  TYPE  SERIES  OF  S.  PRIONOTUS.. 


Iquitos        San  Martin/        Pampa 
Region         VV.  Loreto-       Hermosa 


Rio 
Mishollo 


Tingo 
Maria 


Balta 


Cuzco 
Amazonico 


Explorer's 
Inn 


S.  prionotus 

X 

X 

X 

X 

S.  aculeatus 

X 

X 

S.  fimbriatus 

X 

X 

X 

X 

X 

S.  roseiventris 

X 

X 

^  The  only  other  species  of  Stenocercus  known  from  primarily  the  lowlands  and  lower  Andean  foothills  in 
this  region  is  S.  crassicaiidatus,  which  lacks  a  projecting  vertebral  crest  and  has  a  very  spiny  tail. 

^  The  distributions  of  S.  prionotus  and  S.  aculeatus  overlap  both  altitudinally  and  latitudinally  in  northern 
Peru  but  they  have  not  been  taken  together  at  the  same  locality.  See  Figure  2  for  localities. 

^  In  Manu  National  Park,  S.  fimbriattis  and  S.  roseiventris  are  sympatric  at  Pakdtza,  whereas  S.  prionotus  is 
known  only  from  Cocha  Cashu. 


lizards  obtained  by  Dixon  and  Soini  (1986) 
and  additional  collections  froin  the  region 
made  by  Harvey  Bassler  and  deposited  in 
the  AMNH.  Similarly  large  collections  re- 
sulted from  the  efforts  of  Cadle  and 
McDiarmid  in  Amazonas  (specimens  in 
the  MVZ  and  USNM).  Thus,  barring  ar- 
tifacts introduced  by  the  difficulties  of  col- 
lecting cryptic  rainforest  lizards,  S.  prion- 
otus seeiTis  to  be  absent  froixi  these  sites. 

Documenting  and  explaining  patterns  of 
absence  is  always  difficult,  but  the  appar- 
ent absence  of  Stenocercus  prionotus  froin 
the  lowlands  distant  from  the  Andes  in 
Peru  is  not  due  to  failure  to  collect  Sten- 
ocercus at  these  localities  because  at  least 
one  other  species  of  Stenocercus  is  known 
from  each  (Table  2).  However,  no  lowland 
locality  is  known  in  which  inore  than  two 
species  of  the  "Ophryoessoides  group"  of 
Stenocercus  are  syinpatric.  Of  interest  in 
this  connection  is  that  S.  prionotus  has  not 
been  taken  in  the  upper  reaches  of  the  Rio 
Perene  and  its  tributaries  (Junin  Depart- 
ment), although  it  is  known  from  north 
and  south  of  that  region.  Numerous  spec- 
iinens  of  Stenocercus  of  at  least  five  spe- 
cies (S.  hoettgeri,  S.  crassicaudatus,  S.  for- 
mosus,    S.    scapularis,    and   S.    variabilis'^) 


have  been  collected  along  an  elevational 


^  Based  on  examination  of  two  of  the  three  synty- 
pes  of  Stenocercus  variabilis  Boulenger  (BMNH 
1946.8.11.89,  1946.8.11.91)  I  concur  with  Fritts 
(1974:  66)  that  Boulenger  (1901:  553)  erred  in  as- 


cribing these  specimens  to  "Palca,  Bolixda"  rather 
than  Palca  (Junin  Department),  Peru.  One  specimen 
I  collected  near  Palca,  Peru  (MCZ  178166)  is  nearly 
identical  with  BMNH  1946.8.11.91  in  scale  counts, 
pattern,  and  qualitative  characters  (both  of  these 
specimens  differ  considerably  from  BMNH 
1946.8.11.89  in  color  pattern).  However,  the  attri- 
bution of  the  specimens  to  Peru  is  not  without  some 
equivocation.  The  types  were  collected  by  P.  O.  Si- 
mons (see  Cadle,  1998:  footnote  6),  who  collected  in 
the  vicinity  of  Palca,  Peru,  in  March  and  April,  1900, 
although  that  specific  locality  is  not  listed  in  his  pub- 
lished itinerary  (Chubb,  1919).  However,  Simons's 
itineraiy  places  him  at  "Palca,  18  miles  E  of  La  Paz" 
[Bolivia]  on  9  November  1900  (Chubb,  1919:  5).  Si- 
mons's field  tags  attached  to  the  syntypes  record  sim- 
ply "Palca  3000  m,"  which  is  close  to  the  elevation  of 
Palca,  Peru  (2,740  m),  but  not  that  in  Bolivia  (4,600 
m).  Adding  to  the  confusion  are  entries  in  the 
BMNH  registries  for  the  syntypes,  concerning  which 
Colin  J.  McCarthy  provided  the  following  comments 
via  e-mail: 

Boulenger  originally  wrote  "Palca  Peru  3000m"  but 
later  struck  through  Peru  and  wrote  "Bolivia." 
There  are  two  sheets  of  notes  (presumably  from 
Simons)  stuck  in  at  this  page  of  the  register  about 
the  localities  in  the  batch.  With  regard  to  Palca  he 
has  written  "Palca,  just  S.  of  La  Paz,  Bolivia."  I 
assume  it  was  that  information  that  caused  Boulen- 
ger to  alter  his  original  entry! 

Thus,  the  confusion  may  have  originated  with  the 
note  that  Simons  provided  subsequent  to  cataloguing 
of  the  collection  at  the  BMNH.  In  any  case,  no  spec- 
imens resembling  S.  variabilis  are  definitely  known 
from  Bolivia  and  the  only  species  of  the  genus  defi- 
nitely known  to  occur  above  4,000  m  is  Stenocercus 
chrysopt/gus  from  northern  Peru. 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle 


199 


transect  along  this  well-traveled  route 
(Fritts,  1974;  Cadle,  1991  and  unpublished 
data).  Thus,  analysis  of  circumstantial  dis- 
tributional data  suggests  that  the  distribu- 
tion of  S.  prionotus  in  the  lowlands  of  east- 
ern Peru  may  be  influenced  by  the  num- 
ber of  sympatric  species  of  Stenocercus.  At 
all  localities  from  which  S.  prionotus  has 
been  taken,  only  one  other  species  of  Sten- 
ocercus is  known  (Table  2).  These  patterns 
of  sympatry  are  discussed  in  the  next  sec- 
tion with  reference  to  patterns  of  geo- 
graphic variation  in  S.  prionotus. 

In  contrast,  the  apparent  restriction  of 
Stenocercus  prionotus  to  the  Andean  foot- 
hills and  immediately  adjacent  lowlands  in 
Peru  does  not  seem  related  to  the  distri- 
bution of  any  major  habitat  type  or  phys- 
iographic region.  Most  of  the  extensively 
sampled  localities  (e.g.,  Iquitos  and  Cuzco 
Amazonico)  include  a  variety  of  lowland 
habitats  characteristic  of  western  Amazon- 
ia. Stenocercus  prionotus  is  known  from 
both  floodplain  forests  (Cocha  Cashu)  and 
more  upland  forests  on  river  terraces  in 
southeastern  Peru  (Explorer's  Inn);  Foster 
(1990)  and  Dallmeier  et  al.  (1996)  de- 
scribed these  floristic  communities  in 
southeastern  Peru.  Thus,  the  apparent  ab- 
sence of  S.  prionotus  at  the  localities  dis- 
cussed above  is  not  due  to  some  simple 
relation  to  local  habitat  availability.  For  ex- 
ample, it  is  unclear  why  S.  prionotus  was 
not  obtained  at  Pakitza  (Morales  and 
McDiarmid,  1996),  even  though  it  occurs 
at  nearby  Cocha  Cashu.  On  a  broader  geo- 
graphic scale,  the  restriction  of  S.  priono- 
tus to  lowlands  and  foothills  adjacent  to 
the  Andes  may  be  related  to  the  present 
or  historical  influence  of  the  Andes  on  the 
climate  and  vegetation  (rainfall,  tempera- 
ture, and  major  soil  types)  of  neighboring 
regions. 

A  curious  and  unexplained  hiatus  in  the 
distribution  of  Stenocercus  prionotus  oc- 
curs between  the  vicinity  of  Tingo  Maria— 
Rio  Llullapichis  (Huanuco  Department) 
and  Cocha  Cashu  in  Manu  National  Park 
(Madre  de  Dios  Department),  a  gap  of 
some  600  km  that  includes  the  entire  up- 


per reaches  of  the  Rio  Ucayali-Urubam- 
ba— Ene  system.  Additionally,  populations 
north  and  south  of  this  gap  differ  in  some 
qualitative  and  quantitative  characters  (see 
Patterns  of  Sympatry  and  Geographic  Var- 
iation in  Stenocercus  prionotus,  and  the 
Need  for  Additional  Fieldivork).  Scattered 
collections  (e.g.,  maps  in  Fritts,  1974;  Avi- 
la-Pires,  1995),  but  no  comprehensive  her- 
petofaunal  surveys  or  collections,  are  avail- 
able from  this  vast  region.  Thus,  whether 
the  geographic  hiatus  is  real  or  a  sampling 
artifact  cannot  be  discerned.  Efforts  to  re- 
solve this  issue  need  to  be  made. 

Unconfirmed  Bolivian  Localities  for 
Stenocercus  prionotus.  Because  of  the 
previous  confusion  of  Stenocercus  priono- 
tus wath  S.  caducus,  specimens  perhaps  re- 
ferable to  S.  prionotus  from  several  local- 
ities in  Bolivia  are  unconfirmed.  Boulen- 
ger  (1898)  reported  specimens  of  "Lioce- 
phalus  caducus.  Cope"  in  the  Museo 
Civico  Storia  Naturale  Giacomo  Doria  in 
Genoa  collected  by  Luigi  Balzan  from  four 
localities  (Balzan,  1931).  I  have  not  at- 
tempted to  verify  the  existence  of  these 
specimens.  However,  one  of  Balzan  s  spec- 
imens from  "Barraca,  Rio  Madidi"  was  ex- 
changed to  the  BMNH  (now  BMNH 
98.6.94),  and  is  confirmed  as  S.  prionotus. 
The  "Leiocephalus  caducus"  specimens 
from  the  other  three  Balzan  localities  are 
outside  the  known  distribution  of  S.  cad- 
tictis  but  are  close  to  other  known  localities 
for  S.  prionotus  in  northern  Bolivia  (Fig. 
1).  I  suspect  these  are  S.  prionotus  based 
on  geographic  location.  The  localities  are, 
as  listed  by  Boulenger  (1898)  (see  Fig.  1), 

(1)  "Coroico  and  Chulumani,  Prov.  Yun- 
gas,  1,600  metres  alt."  [La  Paz  Depart- 
ment; Coroico,  1,725  m,  16°10'S,  67°44'W; 
Chulumani,   1,905  m,   16°24'S,  67°31'W]. 

(2)  "Reyes,  right  bank  of  Rio  Beni"  [El 
Beni  Department,  232  m;  14°19'S, 
67°23'W].  (3)  "Missiones  [sic]  Mosetenes" 
[approximately  15°31'S,  67°25'W].5  These 


^  The  Moseten  Indians  inhabited  upper  reaches  of 
the  Rio  Beni  and  its  tributaries  in  the  Andean  foot- 
hills of  the  present  department  of  La  Paz  (Metraux, 


200 


Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


localities  would  not  be  unusual  for  S. 
prionotus,  although  the  first  two  localities 
are  the  highest  elevations  recorded  for  the 
species  (the  species  occurs  at  1500  m  in 
nearby  Puno  Department,  Peru).  All  are 
in  the  upper  reaches  of  the  Rio  Beni, 
whereas  the  two  confirmed  Bolivian  local- 
ities for  S.  prionotus  are  farther  north  in 
the  same  drainage. 

Burt  and  Burt  (1931:  273)  listed  two 
specimens  of  ''Leiocephalus  [  =  Stenocer- 
cus]  scapularis"  (AMNH  22450,  22532) 
from  Rurrenabaque,  Bolivia  [El  Beni  De- 
partment; 14°28'S,  67°34'W].  These  were 
reidentified  in  1971  by  Thomas  H.  Fritts 
as  Stenocercus  caducus  but  they  are  pres- 
ently missing  from  the  AMNH  collection 
(Linda  Ford,  in  litt.,  Februaiy  1999).  Rur- 
renabaque is  very  close  to  Balzan's  locality 
(2)  above.  Based  solely  on  presumed  hab- 
itats and  presently  known  distributions  of 
S.  prionotus  and  S.  caducus  in  eastern  Bo- 
livia (see  text;  Fig.  1),  I  suspect  that 
AMNH  22450  and  22532  are  most  likely 
S.  prionotus.  They  should  be  reexamined 
if  they  are  ever  located.  As  an  outside  pos- 
sibility, any  of  the  unverified  specimens 
from  these  Bolivian  localities  could  rep- 
resent S.  fimbriatus,  which  is  now  known 
from  southern  Peru  (Appendix). 

PATTERNS  OF  SYMPATHY  AND 
GEOGRAPHIC  VARIATION  IN 
STENOCERCUS  PRIONOTUS,  AND  THE 
NEED  FOR  ADDITIONAL  FIELDWORK 

Stenocercus  prionotus  varies  geographi- 
cally in  several  characters,  most  notably  in 
the  height  of  the  vertebral  crest.  Because 


1942).  According  to  Metraux  (1942:  15),  by  the  end 
of  the  19th  century  when  Balzan  collected  his  spec- 
imens the  decimated  population  of  the  Mosetenes 
was  concentrated  in  the  three  "Misiones  Mosetenes" 
of  San  Miguel  de  Muchanes  (15°14'S,  67°39'W),  San- 
ta Ana  (15°31'S,  67°30'W),  and  Covendo  (15°49'S, 
67°06'W).  The  approximate  coordinates  given  in  the 
text  are  the  average  for  these  three  sites.  According 
to  several  maps  with  elevational  contours  indicated, 
Muchanes  and  Covendo  are  between  500  and  1,000 
m  and  Santa  Ana  is  less  than  500  m  in  elevation. 
Mathews  (1879)  also  discusses  these  missions. 


of  the  potential  role  of  the  crest  in  either 
intra-  or  interspecific  communication  it 
seems  appropriate  to  discuss  patterns  of 
geographic  variation  in  S.  prionotus  in  the 
context  of  the  distribution  of  other  species 
of  the  "Ophryoessoides  group."  In  this  re- 
gard, patterns  of  sympatry  among  species 
of  this  group  in  Peru  and  Bolivia  are  es- 
pecially relevant. 

Patterns  of  Sympatry  of  Species  in  the 
"Ophiyoessoides  Group."  The  most  com- 
plex distributional  patterns  for  the 
"Ophryoessoides  group"  are  in  northern 
and  central  Peru,  where  six  species  occur 
(Stenocercus  aculeatus,  S.  fimbriatus,  S. 
prionotus,  S.  huancabambae,  S.  scapularis, 
and  an  undescribed  species;  Fig.  2).  In 
contrast  to  the  other  species,  S.  huanca- 
bambae and  the  undescribed  species  occur 
in  comparatively  dry  deciduous  forests 
west  of  the  known  distributions  for  the 
other  species.  Their  distributions  probably 
do  not  overlap  the  other  species  and  they 
will  not  be  considered  further  here.  Based 
on  known  latitudinal  and  elevational  dis- 
tributions, three  species  (S.  aculeatus,  S. 
fimbriatus,  and  S.  prionotus)  are  probably 
broadly  sympatric  in  northern  Peru,  al- 
though sympatry  is  documented  only  for 
pairs  of  these  at  three  localities  (Fig.  2; 
Table  2).  The  habitat  preferences  of  the 
species  in  the  region  of  sympatry  are  un- 
known. 

Whether  Stenocercus  scapularis  is  sym- 
patric with  S.  prionotus  is  less  clear.  Sten- 
ocercus scapularis  is  known  from  two 
widely  separated  areas  in  Huanuco  and 
Puno  departments  (Peru)  (Fig.  1;  Appen- 
dix), and  it  seems  to  be  elevationally  par- 
apatric  to  S.  prionotus  in  northern  Peru 
(Huanuco  Department).  In  this  area,  S. 
scapularis  occurs  above  1,000  m  and  S. 
prionotus  is  not  known  above  700  m.  In 
Puno  Department,  S.  prionotus  is  known 
from  a  series  obtained  by  Hilda  Heller  at 
San  Juan  del  Oro  (1,520  m  elevation), 
which  elevationally  overlaps  the  distribu- 
tion of  S.  scaptdaris  in  the  same  region 
(1,000-1,830  m).  However,  these  two  spe- 
cies have  not  been  taken  at  the  same  lo- 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle 


201 


cality  in  southern  Peru.  Interestingly,  al- 
though the  largest  series  of  S.  prionotus 
available  from  a  single  locality  (San  Juan 
del  Oro)  is  in  the  elevational  range  of  S. 
scapularis,  no  specimens  of  that  species 
were  obtained  by  Heller  during  a  month- 
long  stay  at  the  site. 

In  southern  Peru  and  Bolivia,  Stenocer- 
cus prionotus  is  not  known  to  be  sympatric 
with  other  members  of  the  "Ophryoesso- 
ides  group."  However,  based  on  the  oc- 
currence of  S.  prionotus  and  S.  finihriatus 
at  two  nearby  localities  within  Manu  Na- 
tional Park  (Cocha  Cashu  and  Pakitza,^  re- 
spectively), sympatry  for  these  two  species 
is  expected  along  the  Andean  front  in  this 
region.  In  southern  Peru  and  Bolivia,  S. 
prionotus  is  also  broadly  sympatric  with  S. 
roseiventris,  a  large  terrestrial  species  that 
is  superficially  similar  to  species  of  the 
"Ophnjoessoides  group."  However,  these 
species  have  been  taken  together  at  only 
one  locality.  Explorer's  Inn. 

To  summarize  these  patterns  of  sympat- 
ly,  the  distributions  of  three  species  of  the 
"Ophryoessoides  group"  overlap  broadly  in 
northern  Peru.  Stenocercus  prionotus  is 
known  to  be  sympatric  with  at  least  one 
other  species  of  the  group  in  Huanuco  and 
Loreto  departments.  No  species  of  the 
"Ophrijoessoides  group"  are  known  to  be 
sympatric  in  southern  Peru  and  Bolivia. 

Geographic  Variation.  Geographic  vari- 
ation pertains  to  both  quantitative  and 
qualitative  characters  in  Stenocercus 
prionotus.  Specimens  from  the  northern 
part  of  the  range  have  higher  average 
counts  of  midbody  scales  than  southern 
specimens,  which  is  shown  graphically  in 
Figure  8.  However,  there  is  broad  overlap 
in  the  counts  from  opposite  ends  of  the 
range.  A  reverse  trend  (lower  counts  in  the 
north),  occurs  in  the  number  of  vertebral 
(dorsal  crest)  scales  (Fig.  9);  this  reflects 
the  reduced  prominence,  and  hence  small- 


''  The  record  of  Stenocercus  fiinbriafiis  from  Pak- 
itza  (Appendix)  extends  the  known  range  for  tliis  spe- 
cies south  by  approximately  200  km  from  Balta  in 
Ucayah  Department,  Peru  (Avila-Pires,  1995). 


er  scales,  of  the  crest  in  southern  popula- 
tions. None  of  these  differences  in  scale 
counts  is  statistically  significant. 

The  height  of  the  vertebral  crest  shows 
strong  clinal  variation  in  Stenocercus 
prionotus.  Populations  from  northern  and 
central  Peru  have  higher  crests  than  those 
from  southern  Peru  and  Bolivia  (Fig.  10). 
However,  the  absence  of  specimens  be- 
tween Huanuco  Department  in  central 
Peru  and  Manu  National  Park  (Madre  de 
Dios  Department)  in  southern  Peru  (Fig. 
1)  makes  it  impossible  to  analyze  this  trend 
in  detail.  Specimens  from  the  former  re- 
gion have  a  high  crest  typical  of  all  speci- 
mens from  that  area  and  farther  north, 
whereas  specimens  from  Madre  de  Dios 
Department  and  farther  south  have  dis- 
tinctly lower  crests.  The  crest  is  not 
strongly  sexually  dimoiphic  in  either  re- 
gion. 

Whether  the  transition  in  crest  height  is 
abrupt  or  gradual  between  central  and 
southern  Peru  is  unclear.  Indeed,  it  is  un- 
clear whether  intei"vening  populations  of 
Stenocercus  prionotus  exist.  Both  patterns 
of  clinal  variation  have  been  extensively 
documented  empirically  and  theoretically 
for  many  organisms  (Endler,  1977).  The 
pattern  of  geographic  variation  in  S.  prion- 
otus could  conform  to  any  of  several  in 
Endler s  (1977:  4)  classification  scheme. 
However,  present  knowledge  of  distribu- 
tions and  character  variation  is  most  simi- 
lar to  Endler's  "differentiated  disjunction" 
pattern,  wherein  disjunct  populations  of  a 
species  diverge  in  one  or  more  characters. 
Further  speculation  is  not  fruitful  in  the 
absence  of  more  extensive  collections  from 
the  region  of  disjunction  in  eastern  Peru. 

A  Hypothesis  of  Causation  for  Geo- 
graphic Differentiation  in  the  Vertebral 
Crest  of  Stenocercus  prionotus.  Variation 
in  the  height  of  the  vertebral  crest  is  a 
prominent  distinction  between  northern 
and  southern  populations  of  Stenocercus 
prionotus.  This  variation  calls  for  an  expla- 
nation while  at  the  same  time  recognizing 
that,  in  the  absence  of  experimental  or 
field   studies,   ascribing  causation  to  pat- 


202         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


48 

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9.3- 
9.6 


11.1- 
11.8 


12.6- 
12.9 


14.2 


14.6- 
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17.5- 
18.3 


20.8- 
21.3 


25.4- 
25.6 


Latitude 


Figure  8.  Geographic  variation  in  the  number  of  midbody  scales  in  Stenocercus  prionotus  (•)  and  S.  caducus  (■).  Each 
symbol  represents  one  specimen;  counts  for  all  specimens  examined  are  plotted.  The  horizontal  axis  represents  degrees  south 
latitude  for  each  locality  converted  to  a  decimal  value  (e.g.,  1  /"SO'  converted  to  1 7.5);  geographically  contiguous  localities  were 
arbitrarily  combined  to  yield  the  latitudinal  ranges  given.  The  S.  caducus  specimens  examined  were  biased  toward  Bolivian 
specimens. 


terns  of  geographic  variation  is  difficult. 
Vertebral  crests  in  iguanids  are  visual  sig- 
nals. The  crests  in  sexually  dimorphic  spe- 
cies are  involved  in  intraspecific  behavioral 
encounters,  especially  intrasexual  aggres- 
sion (e.g.,  Watkins,  1998).  Nonetheless, 
despite  strong  geographic  differentiation, 
the  vertebral  crest  in  Stenocercus  priono- 
tus is  not  strongly  sexually  dimoi*phic  in 
any  part  of  its  range.  Thus,  a  role  in  in- 
trasexual behavioral  signaling  seems  un- 
likely to  be  the  primary  function  of  the 
crest  in  this  species.  Intersexual  signalling 
is  a  possibility,  but  this  would  not  explain 


why  northern   and   southern   populations 
differ  so  strongly  in  this  character. 

Populations  of  Stenocercus  pnonotus 
with  the  highest  vertebral  crests  occur  in 
the  area  of  highest  species  density  and 
documented  sympatry  for  the  ^'Ophrijoes- 
soicles  group"  (Table  2  and  above  discus- 
sion). Not  only  do  northern  populations  of 
S.  prionotus  have  the  highest  vertebral 
crests  within  this  species,  but  in  northern 
and  central  Peru  S.  prionotus  has  a  much 
more  prominent  vertebral  crest  than  other 
species  of  the  "Ophryoessoides  group"  (S. 
aculeatus  approaches  it  most  closely).  In 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle        203 


43 

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6.5- 
7.2 


9.3- 
9.6 


11.1- 
11.8 


12.6- 
12.9 


14.6- 
15.3 


17.5- 
18.3 


20.8- 
21.3 


25.4- 
25.6 


Latitude 


Figure  9. 
axes  are 


Geographic  variation  in  the  number  of  vertebral  crest  scales  in  Stenocercus  prionotus  and  S.  caducus.  Symbols  and 
the  same  as  Figure  8. 


contrast,  in  southern  Peru  and  northern 
BoHvia  the  distribution  of  S.  prionotus 
overlaps  only  S.  scapularis  and  (perhaps) 
S.  fimbriatus  of  the  " Ophrijoessoides 
group,"  but  S.  prionotus  is  not  known  to 
be  sympatric  with  either  of  these  species.' 


'  The  only  case  of  synipatry  of  any  of  these  with 
other  species  of  Stenocercus  of  which  I  am  aware 
involves  S.  prionotus  and  S.  roseiventris  at  Explorer's 
Inn  (cf.  Table  2),  but  S.  roseiventris  is  phenotypically 
dissimilar  to  species  of  the  "Ophryoessoicles  group" 
in  having  shorter  limbs,  different  color  pattern,  spiny 
tail,  and  different  body  form. 


The  vertebral  crest  of  S.  prionotus  in 
southern  Peru  and  northern  Bolivia  is  sim- 
ilar in  development  to  that  in  allopatric 
populations  of  S.  aculeatus. 

I  conjecture  that  the  strongly  developed 
vertebral  crests  of  Stenocercus  prionotus 
in  northern  Peru  may  be  related  to  the 
presence  of  multiple  sympatric  congeners 
in  that  part  of  the  range,  and  that  the  crest 
functions  as  a  species  recognition  charac- 
ter. If  true,  this  explanation  for  variation  in 
crest  height  would  have  no  precedent 
among  lizards.  The  only  analogous  situa- 


204         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


(a)     prionotus 

Huanuco,  Peru 
67  mm 


(b)  prionotus 
Puno,  Peru 
73  mm 


(c) 


prionotus 
Beni,  Bolivia 
81  mm 


(d)     caducus 

Sta.  Cruz,  Bolivia 
72  mm 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle        205 


tion  seems  to  be  dewlaps  in  Anolis,  in 
which  species  recognition  has  been  pos- 
tulated as  a  biological  role  for  dewlap  color 
and  size  in  complex  assemblages  of  these 
lizards  (Williams  and  Rand,  1977;  Losos 
and  Chu,  1998). 

The  Need  for  Additional  Fieldwork. 
Only  field  observations  and  experimental 
studies  of  the  function  of  the  vertebral 
crest  in  Stenocercus  prionotus  can  deter- 
mine whether  the  functional  hypothesis 
advanced  here  is  correct.  Ideally,  parallel 
studies  should  be  conducted  in  the  area  of 
sympatry  with  other  species  of  the 
"Ophryoessoides  group,"  as  well  as  in  the 
area  with  no  known  sympatric  species  of 
that  group.  That  design  would  permit  an 
evaluation  of  any  context-dependent  use  of 
the  crest  in  these  populations.  Two  sym- 
patric species  are  known  from  the  vicinity 
of  Tingo  Maria  (Huanuco  Department; 
Table  2),  a  very  accessible  area  for  study. 

Two  other  reasons  for  additional  com- 
prehensive fieldwork  on  these  species  are 
indicated.  First,  we  need  documentation 
of  the  extent  to  which  species  of  the 
"Ophryoessoides  group"  are  sympatric  or 
syntopic.  Particular  targets  should  be  the 
northern  part  of  the  range  of  Stenocercus 
prionotus,  where  S.  aculeatus  and  S.  fim- 
hriatus  are  known  to  have  overlapping 
ranges;  and  Manu  National  Park  in  south- 
ern Peru,  a  reasonably  accessible  area 
where  S.  jimbriatus  and  S.  prionotus  are 
known  to  be  closely  allopatric  (but  not 
sympatric)  on  a  microgeographic  scale  (Ta- 
ble 2).  Second,  the  character  differences 
between  the  northern  populations  of  S. 
prionotus  and  those  of  southern  Peru  and 
Bolivia  could  indicate  that  two  or  more 
species  reside  within  my  concept  of  this 
species.    Fieldwork    concentrated    in    the 


geographic  gap  between  Tingo  Maria  and 
vicinity  (Huanuco  Department)  and  Cocha 
Cashu  (Madre  de  Dios  department),  the 
two  most  proximate  localities  of  the  north- 
ern and  southern  population  groups,  is 
needed  to  verify  whether  intermediate 
populations  occur.  The  possibility  that 
these  might  be  two  allopatric  species  sep- 
arated by  a  broad  geographic  gap  should 
not  be  dismissed  without  further  evalua- 
tion. 

NATURAL  HISTORY  OF  STENOCERCUS 
PRIONOTUS 

Few  natural  history  observations  exist 
for  Stenocercus  prionotus.  Wade  C.  Sher- 
brooke  (field  notes)  found  the  holotype 
running  in  the  relatively  open  floor  of  a 
bamboo  garden  at  about  1200  h.  USNM 
193685  was  taken  along  a  trail  within  rain- 
forest. Brief  notes  associated  with  the  se- 
ries from  Explorers  Inn  (Madre  de  Dios 
Department,  Peru;  USNM  247468-69, 
247680,  269022)  indicate  that  specimens 
were  obtained  during  the  day  on  the 
ground  from  clearings  around  the  lodge, 
although  USNM  269022,  an  adult  female 
(91  mm  SVL),  was  on  a  leaf  40  cm  above 
the  ground.  MCZ  150243  was  retrieved 
from  a  mist  net  near  the  edge  of  tall  flood- 
plain  forest  at  Cocha  Cashu.  In  the  dry 
season  (September— December)  of  a  low- 
land rainforest  in  the  vicinity  of  Tumi  Chu- 
cua  (Beni  Department,  Bolivia),  Fugler 
(1986:  table  5)  found  Stenocercus  priono- 
tus (reported  as  Ophryoessoides  acideatus) 
in  varzea  (seasonally  inundated  rainforest) 
but  not  in  terra  firme  rainforests  of  the 
area;  females  with  enlarged  eggs  were 
found  in  early  November  (Fugler,  1986: 
table  4). 

Hilda  H.  Heller  provided  the  foUov^dng 


Figure  10.  Diagrammatic  representation  of  geographic  and  size-related  variation  in  the  height  of  the  vertebral  crest  in  Steno- 
cercus prionotus  and  S.  caducus.  Drawings  were  made  with  a  camera  lucida  to  emphasize  the  form  and  height  of  the  vertebral 
crest.  Sketches  are  drawn  to  an  approximately  uniform  interval  between  the  external  ear  opening  (EE)  and  the  white  antehumeral 
stripe  (AS).  For  each  specimen  the  geographic  location  and  the  SVL  are  given  (all  specimens  are  adult  males):  (a)  MCZ  43759, 
(b)  FMNH  64799,  (c)  USNM  280250,  (d)  CM  970.  Note  especially  the  differences  between  the  northern  specimen  of  S.  prionotus 
(a)  compared  to  southern  ones  (b  and  c),  especially  given  the  size  differences  among  these;  and  the  differences  between  size- 
matched  specimens  of  S.  prionotus  and  S.  caducus  (b  and  d). 


206         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


notes  on  the  San  Juan  [del  Oro]  locality 
from  which  she  obtained  a  series  of  Sten- 
ocercus  prionotus  in  the  early  1950s  (K.  P. 
Schmidt  archives,  FMNH): 

Steep  forest  with  deep  undergrowth.  Steep  fields. 
Rainfall  probably  somewhat  greater  than  at  Pampa 
Grande,^  due  to  its  colder  climate  and  steep  ex- 
posure; I  have  no  figures.  Brushy  second  growth 
may  be  burned  in  fairly  wide  patches  in  August, 
and  [the]  resort  is  frequently  made  to  burning,  in- 
choating a  moderately  dry  winter  period. 

As  of  the  mid-1980s  very  little  undisturbed 
forest  was  left  in  the  vicinity  of  San  Juan 
del  Oro  (personal  observations).  Although 
Heller  provided  detailed  notes  on  some  of 
the  snakes  and  frogs  from  her  collection, 
she  makes  no  specific  comments  about  the 
lizards. 

Most  observations  suggest  that  Steno- 
cercus  prionotus  prefers  open  habitats, 
such  as  areas  of  human  disturbance  and 
light  gaps  within  forests  (e.g.,  created  by 
trails),  rather  than  deep  rainforests.  Alter- 
natively, the  observations  may  simply  in- 
dicate the  ease  of  observation  and  capture 
in  more  open  habitats.  A  combination  of 
cryptic  coloration  and  escape  behavior 
(rapid  flight  followed  by  immobility)  pos- 
sibly makes  S.  prionotus  very  difficult  to 
observe  in  closed-canopy  rainforest,  as  re- 
ported for  the  similar  species,  S.  fimbria- 
tus  (Dixon  and  Soini,  1986;  Avila-Pires, 
1995)  and  S.  caducus  (Scrocchi  et  al., 
1985).^  However,  we  currently  lack  obser- 
vations to  support  these  statements  for  S. 
prionotus. 

COMPARISON  OF  STENOCERCUS 
PRIONOTUS  WITH  S.  CADUCUS 

A  scaly  flap  associated  with  the  posthu- 
meral  pockets  is  a  unique  and  unquestion- 
ably derived  character  shared  by  Stenocer- 
cus  prionotus  and  S.  caducus,  which  sug- 
gests that  these  are  sister  species  (Figs.  5, 


11).^"  I  am  unaware  of  a  similar  structure 
in  any  other  lizards.  Some  individual  vari- 
ation in  the  precise  form  and  size  of  the 
flap  occurs  in  both  species  but  it  seems 
extremely  iinprobable  that  these  structures 
are  not  homologous  in  the  two  species. 

Moreoever,  Stenocercus  prionotus  and 
S.  caducus  are  similar  in  standard  meristic 
characters  (Table  1;  Figs.  8,  9),  and  the 
siinilarity  among  Bolivian  populations  of 
both  species  has  caused  confusion  about 
the  identity  of  particular  populations  (see 
citations  in  the  synonymy  of  S.  prionotus) . 
Geographic,  ontogenetic,  or  individual  var- 
iation of  some  characters  within  both  spe- 
cies, especially  the  height  of  the  vertebral 
crest  and  the  number  of  midbody  scale 
rows  (Fig.  8),  further  clouds  the  distinc- 
tions between  them.  Differentiating  the 
northern  populations  of  S.  prionotus  from 
S.  caducus  is  unequivocal  and  facile  based 
solely  on  the  size  of  the  vertebral  crest  and 
on  associated  meristic  counts.  However, 
specimens  of  S.  prionotus  from  southern 
Peru  and  northern  Bolivia  are  more  diffi- 
cult to  distinguish  from  S.  caducus.  For 
example,  animals  from  populations  of  S. 
prionotus  in  southern  Peru  and  Bolivia 
have  less  prominent  vertebral  crests 
(hence,  higher  vertebral  scale  counts),  and 
generally  lower  numbers  of  midbody  scale 
rows,  than  do  specimens  from  northern 
populations  (Figs.  8,  9);  in  these  respects 
they  are  more  similar  to  S.  caducus.  Nev- 
ertheless, even  accounting  for  these  diffi- 
culties, a  combination  of  three  qualitative 
characters  is  sufficient  to  distinguish  S. 
prionotus  from  S.  caducus,  and  the  species 


^  I  have  been  unable  to  localize  Pampa  Grande. 

®  Cei  (1993)  claimed  that  Stenocercus  caducus  was 
arboreal,  but  Scrocchi  et  al.  (1985)  reported  the  be- 
havior of  this  species  in  more  detail  and  stated  that 
it  was  terrestrial. 


'"  In  my  comparisons  I  have  emphasized  Bolivian 
specimens  referred  to  Stenocercus  caducus,  whereas 
the  type  locality  is  "Paraguay."  I  have  not  fully  con- 
vinced myself  that  specimens  referred  to  this  species 
from  Bolivia,  Paraguay,  and  Argentina  are,  in  fact,  all 
the  same  taxon.  Considerable  variation  exists  in  some 
aspects  of  coloration  and  scale  characters  in  diese 
specimens.  However,  my  concept  of  S.  caducus  cor- 
responds to  that  used  in  current  literature  (e.g.,  Gal- 
lardo,  1959;  Scrocchi  et  al,  1985;  Cei,  1993).  Only  a 
thorough  study  of  S.  caducus  across  its  range  will  re- 
solve this  issue. 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Codle        207 


may  differ  in  patterns  of  sexual  size  di- 
morphism.^^ 

The  Form  of  the  Vertebral  Crest.  De- 
spite geographic  variation  in  the  promi- 
nence of  the  vertebral  crest,  males  and  fe- 
males of  Stenocercus  prionotiis  have  a  dis- 
tinctly projecting  serrate  vertebral  crest  ex- 
tending from  the  nuchal  region  to  the 
anterior  portion  of  the  tail  (Figs.  6,  10,  12). 
The  scales  of  the  crest  are  strongly  trian- 
gular in  lateral  view,  are  flaplike  (i.e.,  they 
bend  easily),  project  vertically  from  the 
dorsum,  and  are  strongly  differentiated 
from  the  adjacent  dorsal  scales.  Although 
the  crest  is  somewhat  less  developed  in  fe- 
males, it  is  prominent  in  both  sexes.  Spec- 
imens from  northern  Bolivia  and  southern 
Peru  have  a  substantially  lower  crest  than 
specimens  from  central  and  northern  Peru 
(Fig.  10).  Nonetheless,  the  form  and  pro- 
jection angle  of  the  crest  scales  is  the  same 
as  in  the  northern  populations. 

In  contrast,  the  scales  of  the  vertebral 
crest  in  Stenocercus  caducus  are  only  mod- 
erately differentiated  from  adjacent  dorsal 
scales  in  being  more  strongly  keeled  and 
mucronate.  The  crest  in  S.  caducus  is  only 
slightly  projecting  in  males  (Fig.  10)  and 
even  less  so  in  females  (Fig.  12);  the  crest 
is  mainly  apparent  on  the  neck  and  ante- 
rior body.  In  S.  caducus,  the  scales  of  the 
crest  are  stiff  and  prismatic,  and  the  main 
axis  of  projection  is  posterior  rather  than 
vertical,  as  in  S.  prionotiis. 

Crest  height  in  Stenocercus  varies  posi- 
tively with  size  and  thus  it  is  critical  to 
compare  similar-sized  specimens  when 
documenting  differences  among  popula- 
tion samples  or  species.  This  realization 
has  been  critical  to  differentiating  Steno- 
cercus prionotiis  from  S.  caducus  in  south- 


"  I  am  uncertain  how  Fugler  (1983,  1986,  1989) 
distinguished  Boh\dan  specimens  he  referred  to  Sten- 
ocercus aculeatus  and  S.  caducus.  In  1983  and  1986 
he  referred  specimens  from  Tumi  Chucua  (Beni,  Bo- 
livia) to  S.  aculeatus.  In  1989  he  Usted  these  again, 
along  with  ROM  specimens  from  San  Marcos  Ranch 
(Beni,  Bolivia)  identified  as  S.  caducus.  Fugler  spec- 
imens from  these  localities  that  I  have  examined  are 
all  S.  prionotus  (see  list  of  paratypes). 


Figure  1 1 .  Axillary  region  of  Stenocercus  caducus  showing 
the  posthumeral  flap  (MCZ  34215).  Anterior  to  the  left.  The 
posthumeral  flap  comprises  the  heavily  outlined  scales  pos- 
teroventral  to  the  forelimb.  The  opening  of  the  posthumeral 
pocket  is  the  heavily  stippled  area  deep  to  the  flap.  Top,  Pos- 
thumeral flap  in  its  normal  orientation  covering  the  anterov- 
entral  portion  of  the  pocket.  Bottom,  the  flap  deflected  ven- 
trally,  with  its  posterior  scales  viewed  from  their  tips.  Approx- 
imately X8.5. 


ern  Peru  and  Bolivia.  Large  adult  males  of 
S.  prionotus  from  southern  populations 
are  scarce  in  collections.  For  example,  al- 
though only  18  specimens  of  S.  prionotus 
are  available  from  northern  Peru,  one  half 
of  these  are  males  with  SVL  >60  mm.  In 


208         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


contrast,  twice  as  many  specimens  each  of 
S.  prionotus  and  S.  caducus  are  available 
from  southern  Peru  and  Bolivia.  Yet,  only 
one  third  of  the  available  specimens  of  ei- 
ther species  from  these  areas  are  males 
>60  mm  SVL,  and  no  males  of  S.  caducus 
were  >72  mm  SVL. 

Figure  10  shows  differences  in  the 
height  of  the  vertebral  crest  in  a  series  of 
males  of  Stenocercus  prionotus  from 
southern  Peru  and  Bolivia  compared  with 
similar-sized  specimens  of  S.  prionotus 
from  northern  Peru  and  with  S.  caducus 
(the  largest  males  of  S.  caducus  studied 
were  72  mm  SVL;  see  also  Fig.  6).  The 
trend  toward  lower  crests  in  S.  prionotus 
from  the  southern  part  of  its  range  is  evi- 
dent, as  is  the  difference  between  S.  cad- 
ucus and  S.  prionotus.  A  comparison  of  all 
specimens  suggests  that  the  difference  in 
the  height  of  the  vertebral  crest  between 
S.  prionotus  and  S.  caducus  males  begins 
to  be  apparent  by  approximately  65  mm 
SVL  and  becomes  pronounced  at  around 
70-75  mm  SVL.  No  males  of  S.  caducus 
>72  mm  SVL  were  among  the  specimens 
examined,  despite  the  availability  of  a  large 
number  of  specimens  froin  southern  Bo- 
livia, including  a  series  of  31  specimens 
(10  males  >60  inm  SVL)  from  the  vicinity 
of  Santa  Cruz.  A  siniilar  contrast  in  crest 
height  appears  in  females  of  the  two  spe- 
cies (Fig.  12). 

Angulate  Temporal  Scales.  Stenocercus 
prionotus  has  two  (occasionally  three)  very 
strongly  keeled,  projecting  angulate  tein- 
poral  scales  on  each  side  (Fig.  4).  These 
are  much  larger  than  adjacent  scales  on 
the  head  and  they  are  partially  or  com- 
pletely separated  from  the  large  posterior 
head  scales  (parietals,  postparietals,  and 
occipital)  by  one  row  of  small  keeled  scales 
(occasionally  partially  doubled).  Stenocer- 
cus caducus  usually  has  two  (occasionally 
three)  angulate  temporals  that  are  smaller 
and  less  projecting  than  those  in  S.  prion- 
otus. In  S.  caducus  the  angulate  temporals 
may  or  may  not  be  larger  than  adjacent 
posterior   head   scales    and   they   are   not 


Figure  12.  Size-matched  females  of  Stenocercus  prionotus 
and  S.  caducus  from  Bolivia.  Top,  S.  prionotus  (USNM 
269022,  snout-vent  length  [SVL]  91  mm).  Bottom,  S.  caducus 
(UTA  38046,  SVL  93  mm).  Note  the  subtle  difference  in  crest 
height  between  the  two  specimens  and  their  otherwise  similar 
patterns. 


strongly  differentiated  from  other  posteri- 
or head  scales. 

Color  Pattern  of  the  Gular  Region. 
Many  specimens  of  Stenocercus  prionotus 
have  a  regular  pattern  of  alternating  diag- 
onal light  and  dark  stripes  on  the  throat. 
These  usually  converge  closely  toward  the 
midline  (Fig.  7)  and  are  most  easily  visu- 
alized in  preseived  specimens  submerged 
in  alcohol.  This  pattern  consists  of  a  dark 
stripe  beginning  at  a  point  on  the  lower 
labials  in  line  with,  but  broader  than,  the 
subocular  dark  bar.  The  stripe  projects 
posteromedially,  gradually  fading  and 
blending  with  the  ventral  ground  color  on 
the  neck  anterior  to  the  pectoral  region. 
The  dark  stripe  is  bordered  on  either  side 
by  a  distinct  pale  stripe.  Anteriorly,  this  se- 
ries is  preceded  by  another  dark  and  an- 
other pale  stripe.  The  dark  stripes  are  usu- 
ally approximately  twice  as  wide  as  the 
pale  ones,  although  not  always  (e.g.,  the 
dark  stripes  are  only  slightly  wider  than 
the  pale  ones  in  MCZ  150243).  In  life  the 
pattern  may  manifest  itself  as  a  series  of 
pale  stripes  on  a  darker  background  (e.g., 
the  "gular  area  streaked  by  several  light 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle 


209 


cream  colored  lines"  in  the  life  colors  of 
the  holotype). 

The  gular  region  appears  uniform  in 
many  preserved  specimens  of  Stenocercus 
prionotus,  but  I  suspect  this  is  a  preser- 
vation artifact.  Occasional  specimens  have 
pale  spots  in  the  pectoral  region,  and  oth- 
ers are  essentially  unicolor  and  without  ap- 
parent pattern  (again,  probably  a  preser- 
vation artifact). 

On  the  other  hand,  the  throat  pattern  of 
Stenocercus  caducus  is  highly  variable  and 
irregular.  When  a  distinctive  pattern  is 
present,  it  most  often  consists  of  light 
spots  rather  than  alternating  stripes  (Fig. 
13).  Cope  (1862)  described  the  holotype 
of  S.  caducus  from  Paraguay  as  having  a 
dark  throat  that  was  "light  varied"  (i.e., 
variegated,  or  spotted),  and  some  speci- 
mens I  examined  have  this  pattern  (Fig. 
13).  None  of  several  color  descriptions  for 
Argentinian  specimens  of  S.  caducus  men- 
tion stripes  or  spots  on  the  throat.  Scrocchi 
et  al.  (1985)  described  living  examples  as 
having  pale  spots  in  parallel  transverse 
rows  in  the  pectoral  region  or  with  pale 
spots  on  the  abdomen,  but  did  not  com- 
ment on  the  throat  pattern;  Gallardo 
(1959)  described  the  ventral  coloration  as 
"pale  olive  with  some  scattered  pale  spots; 
throat  darker";  and  Cei  (1993)  described 
the  venter  as  "dark  brownish  with  series  of 
rounded  pale  spots,  sometimes  anastomos- 
ing along  the  length  of  a  median  line."  Al- 
though no  authors  mention  alternating 
light  and  dark  stripes  on  the  throat  in  S. 
caducus,  UTA  38046  does  have  this  pat- 
tern (Fig.  13).  But  in  this  specimen  the 
stripes  are  confined  to  the  lateral  edges  of 
the  throat  (i.e.,  do  not  closely  approach  the 
midline  as  in  S.  prionotus).  Apart  from  the 
throat  pattern,  the  coloration  of  S.  prion- 
otus and  S.  caducus  seems  to  be  veiy  sim- 
ilar judging  from  descriptions  of  S.  cadu- 
cus in  the  literature  (Gallardo,  1959; 
Scrocchi  et  al,  1985;  Cei,  1993). 

A  Possible  Dijference  in  Sexual  Size  Di- 
morphism. Data  presented  in  Table  1  sug- 
gests another  contrast  between  Stenocer- 
cus prionotus  and  S.  caducus:  S.  prionotus 


Figure  13.  Gular  patterns  in  Stenocercus  caducus.  Top,  typ- 
ical throat  pattern  consisting  of  light  spots  on  a  dark  back- 
ground (BMNH  1927.8.1.163).  Bottom,  variant  pattern  consist- 
ing of  stripes  confined  to  the  lateral  portion  of  the  throat  (UTA 
38046).  Connpare  to  Figure  7. 


210         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


is  not  dimorphic  in  the  maximum  sizes  at- 
tained by  males  and  females,  whereas  fe- 
males of  S.  caducus  apparently  attain 
about  20  mm  greater  SVL  than  males. 
However,  estimating  maximum  size  is  sub- 
ject to  considerable  sampling  error  so  this 
distinction  should  be  considered  to  be  only 
provisional.  Nonetheless,  males  of  S. 
prionotus  with  SVL  >80  mm  are  known 
from  the  northern  (USNM  193685)  and 
southern  (USNM  280250,  BMNH 
98.6.9.4)  portions  of  its  range,  even  though 
the  three  largest  males  from  the  largest 
population  sample  (23  specimens  in 
FMNH  from  Puno  Department,  Peru) 
had  an  SVL  of  73  mm  (this  sample  includ- 
ed six  adult  females  with  an  SVL  of  78-89 
mm). 

In  contrast,  a  sample  of  39  Stenocercus 
caducus  from  Bolivia  included  12  adult 
males,  none  of  which  had  an  SVL  >72 
mm;  in  the  total  sample  of  S.  caducus  iJSJ 
=  43)  17  females  had  an  SVL  >80  mm 
(range  80-93  mm).^-  Thus,  unless  a  sys- 
tematic collecting  bias  against  males  exists, 
the  different  pattern  of  sexual  size  dimor- 
phism provides  another  character  distin- 
guishing S.  prionotus  and  S.  caducus.  Data 
presented  in  Table  1  suggests  that  other 
species  of  the  " Ophrijoessoides  group" 
may  be  size  dimoi*phic  (S.  fimbriatus,  S. 
hiiancabambae,  and  ?S.  scapularis)  or  not 
(S.  aculeatus),  and  either  males  (S.  huan- 
cahambae)  or  females  (S.  caducus  and  S. 
fimbriatus)  may  attain  a  larger  body  size. 

DISTRIBUTIONS  OF  STENOCERCUS 
PRIONOTUS  AND  S.  CADUCUS  IN 
EASTERN  BOLIVIA 

The    ranges    of   Stenocercus   prionotus 
and  S.  caducus  approach  one  another  in 


eastern  Bolivia,  but  do  not  overlap. ^^  Cur- 
rently, the  two  closest  documented  locali- 
ties are,  for  S.  prionotus,  near  the  junction 
of  the  Rio  Madidi  and  the  Rio  Beni 
(BMNH  98.6.9.4),  and  for  S.  caducus, 
many  specimens  from  the  vicinity  of  Santa 
Cruz  de  la  Sierra  (see  above  discussion  for 
S.  prionotus  and  Appendix;  Fig.  1).  Sten- 
ocercus caducus  is  also  known  from  the 
Bolivia— Brazil  border  in  the  region  of  the 
Serrania  de  Huanchaca  in  northern  Santa 
Cruz  Department,  Bolivia. ^^ 

I  am  aware  of  no  specimens  of  Steno- 
cercus referable  to  either  S.  caducus  or  S. 
prionotus  between  the  Rio  Beni  valley  and 
roughly  a  line  connecting  Santa  Cnaz  and 
the  vicinity  of  Serrania  de  Huanchaca. 
Southeast  of  the  Rio  Beni,  the  central  part 
of  the  Beni  basin  (the  Llanos  de  Mojos)  is 
characterized  by  flooded  savanna  grass- 
lands, palm  savannas,  swamps,  and  other 
habitats  that  are  inundated  for  significant 
portions  of  the  year;  terra  firme  forests  are 
restricted  to  somew^hat  elevated  levees 
along  rivers  (Clapperton,  1993:  196;  Han- 
agarth,  1993).  Stenocercus  prionotus  or  S. 
caducus  seem  unlikely  to  occur  in  this  area 
except  possibly  in  these  galleiy  forests,  al- 
though Fugler  (1986)  reported  S.  priono- 
tus (as  Ophryoessoides  aculeatus)  in  sea- 
sonally flooded  forest  during  the  dry  sea- 
son. 

Stenocercus  prionotus  and  S.  caducus 
probably  are  segregated  by  habitat  in  Bo- 
livia and  their  distributions  may  not  over- 
lap.   Stenocercus   prionotus    is    associated 


^-  These  sizes  are  somewhat  larger  than  those  pre- 
viously reported  (81  mm;  Gallardo,  1959;  Scrocchi  et 
al,  1985;  Marcus,  1986).  Cei  (1993)  stated  that  S. 
caducus  reaches  only  75  mm  SVL  in  Argentina.  Sexes 
were  not  given  for  any  individual  or  sexed  specimens 
in  these  reports. 


'"^  All  references  to  "Ophryoessoides  aculeatus"  in 
Bolivia  (e.g.,  Fugler,  1983,  1986)  that  I  verified  have 
referred  to  Stenocercus  prionotus.  However,  given 
the  general  confusion  of  species  in  this  complex, 
some  records  not  traced  will  have  to  be  checked  to 
rule  out  the  possibility  that  they  do  not  refer  to  S. 
caducus  or  perhaps  some  other  species  of  the 
"Ophryoessoides  group,"  such  as  S.  fimbriatus  or  S. 
scapidaris  (see  Distribution  Patterns  in  Stenocercus 
prionotus). 

i-i  See  the  Appendix,  UTA  38048.  Michael  Hawey 
(personal  communication)  recently  obtained  speci- 
mens of  Stenocercus  caducus  at  El  Refugio,  a  lowland 
locality  at  the  southern  end  of  the  Serrania  de  Huan- 
chaca (14°44'S.  61°01'W). 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle 


211 


with  upper  Amazonian  and  lower  montane 
rainforests  with  annual  rainfall  greater 
than  2,000  mm  in  both  Peru  and  Bolivia. 
On  the  other  hand,  confirmed  localities  of 
S.  caducus  are  within  the  physiographic 
domain  broadly  referred  to  as  chaco,  in- 
cluding a  mixture  of  diy  forests,  palm  sa- 
vannas, galleiy  forests,  deciduous  forests, 
and  ecotonal  areas  (Scrocchi  et  al.,  1985; 
Marcus,  1986;  Cei,  1993).  Short  (1975) 
and  Parker  et  al.  (1993)  described  the  di- 
versity of  chaco  habitats.  Gallardo  (1979: 
table  12.1)  listed  S.  caducus  as  a  species 
"basically  restricted  to  the  chaco."  Average 
annual  rainfall  in  this  area  is  less  than 
1,000  mm.  Stenocercus  caclucus  is  known 
from  Parque  Nacional  Noel  Kempff  Mer- 
cado  and  vicinity  in  Bolivia  (see  footnote 
14;  Haivey,  1998).  This  area  is  character- 
ized by  a  complex  mixture  of  habitat  types, 
including  deciduous  forests  and  cerrado 
enclaves,  and  with  an  annual  rainfall  of 
1,400-1,500  mm  (Killeen,  1998).  Hai-vey 
(1998)  encountered  S.  caducus  at  granitic 
outcrops  covered  by  semideciduous  forests 
and  more  open  habitats.  The  herpetofauna 
of  this  site  is  a  mixture  of  species  that  are 
typical  of  Amazonian  and  of  chaco  envi- 
ronments (Hai^vey,  1998;  personal  obser- 
vations). 

The  range  of  Stenocercus  caducus  ex- 
tends outside  the  strictly  defined  chaco  re- 
gion (see  Short,  1975,  and  Clapperton 
1993,  for  discussion)  on  the  southeastern 
edge  of  its  range  east  of  the  Rio  Paraguay 
and  in  the  Andean  foothills  of  southern 
Bolivia  and  northern  Argentina  (Fig.  1). 
Harvey  (1997)  reported  S.  caducus  from 
"subtropical  wet  forests"  (1,150—2,050  m 
elevation)  in  southern  Bolivia.  He  charac- 
terized S.  caducus  as  a  "Chacoan  species 
that  invade[s]  the  Andean  foothills  .  .  .  [in- 
cluding] those  distributed  within  the  Gran 
Chaco  or  that  occur  in  diy  forests  sur- 
rounding the  Gran  Chaco"  (Hai-vey,  1997: 
35).  The  montane  wet  forests  (yungas)  of 
this  area  are  restricted  to  ridges  high 
enough  for  cloud  formation  during  much 
of  the  year  (generally  >  1,500  m  elevation), 
and  they  are  surrounded  by  deciduous  dry 


forest  (Schulenberg  et  al.,  1997).  The  cli- 
mate of  this  area  is  generally  dry  and  it 
receives  only  about  1,200  mm  of  rainfall 
per  year  (Hoist,  1997). 

The  transition  between  the  wet  rainfo- 
rests of  Peru  and  northern  Bolivia  (range 
of  Stenocercus  prionotus)  and  the  chaco 
habitats  (range  of  S.  caducus)  occurs  in  a 
very  broad  ecotone  consisting  of  savannas, 
evergreen  shrublands,  and  gallery  forests 
of  the  Beni  basin  and  Rio  Mamore  drain- 
age, from  which  no  specimens  of  either  S. 
prionotus  or  S.  caducus  have  been  report- 
ed. The  piedmont  forests  of  the  Andes  be- 
tween the  known  ranges  of  S.  prionotus 
and  S.  caducus,  which  are  wetter  than  ad- 
jacent lowland  forests  because  of  the  mod- 
erating effect  of  the  Andes,  provide  one 
potential  route  for  contact  or  overlap  of 
their  ranges. 

The  eastern  distributional  limits  of  Sten- 
ocercus caducus  along  the  Bolivia— Brazil 
frontier  are  not  well  understood.  I  am  un- 
aware of  verified  records  from  Brazil,  al- 
though the  species  does  occur  close  to  the 
Brazilian  border  in  the  vicinity  of  the  Ser- 
rania  de  Huanchaca  in  Parque  Nacional 
Noel  Kempff  Mercado.  Some  references 
to  "Stenocercus  caducus"  from  western 
Brazil  (e.g.,  Mato  Grosso  State;  Cope, 
1887;  Boulenger,  1903)  likely  refer  instead 
to  an  undescribed  species  veiy  similar  to 
S.  caducus  (P.  E.  Vanzolini  and  E.  E.  Wil- 
liams, personal  communication;  personal 
observations).  However,  the  ranges  of  S. 
caducus  and  the  undescribed  species  in 
eastern  Bolivia-Paraguay  and  western  Bra- 
zil are  not  well  defined;  the  two  species 
may  be  separated  by  the  seasonally  inun- 
dated savannas  of  the  pantanal.  Addition- 
ally, few  specimens  of  S.  caducus  appar- 
ently exist  from  the  chaco  of  northwestern 
Paraguay,  although  Aquino  et  al.  (1996)  re- 
ported specimens  from  Parque  Nacional 
Defensores  del  Chaco  (approximately 
20°30'S,  60°20'W),  as  well  as  other  Para- 
guayan localities  in  more  mesic  regions 
east  of  the  Rio  Paraguay. 


212         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


IS  THE  DISTRIBUTION  OF 
STENOCERCUS  ACULEATUS 
DISJUNCT? 

In  the  process  of  diagnosing  Stenocercus 
prionotus  I  reviewed  the  characters  and 
distribution  of  S.  aculeatus.  In  addition  to 
variation  in  some  characters  of  uncertain 
significance,  some  aspects  of  the  distribu- 
tion of  S.  aculeatus  seem  pecuHar  (Fig.  2). 
First,  the  distribution  of  S.  aculeatus  ap- 
pears to  be  disjunct  between  northern 
Peru  and  eastern  Ecuador.  Although  the 
type  locality  is  in  northern  Peru  (Moya- 
bamba,  San  Martin  Department),  most 
specimens  are  from  eastern  Ecuador  (Fig. 
2).  The  two  areas  from  which  specimens 
are  known  (Fig.  2)  are  separated  by  a 
broad  geographic  gap  through  which 
courses  the  main  tributary  of  the  upper 
Amazon,  the  Rio  Marafion.  Neither  S.  acu- 
leatus nor  any  other  species  of  Stenocercus 
was  obtained  during  hei"petofaunal  surveys 
in  northern  Loreto  Department,  Peru 
(Duellman  and  Mendelson,  1995),  north- 
ern Amazonas  Department,  Peru  (Rio  Ce- 
nepa  and  Rio  Santiago;  J.  E.  Cadle  and  R. 
W.  McDiarmid,  unpublished  data),  or  dur- 
ing a  rapid  biological  assessment  of  the 
Cordillera  del  Condor  region  of  southeast- 
ern Ecuador  and  northern  Peru  (Schulen- 
berg  and  Awbrey,  1997a).  Stenocercus  acu- 
leatus is  known  from  many  localities  in  ad- 
jacent regions  of  Ecuador. 

Second,  all  Ecuadorian  localities  for 
Stenocercus  aculeatus  are  in  the  drainages 
of  the  Rio  Pastaza  and  the  Rio  Curaray. 
No  specimens  are  known  from  the  Rio 
Napo  drainage  just  to  the  north,  even 
though  no  recognized  physiographic  or 
faunal  break  seems  to  separate  the  Rio 
Napo  drainage  from  the  Rio  Curaray 
drainage.  However,  all  localities  that  have 
been  sampled  comprehensively  from  the 
Rio  Napo  are  on  the  left  (northern)  bank 
of  the  river  (e.g.,  Duellman,  1978;  Vitt  and 
De  la  Torre,  1996;  unpublished  list  from  a 
large  collection  from  the  Jatun  Sacha  Bi- 
ological Station  assembled  and  under 
study  by  Gregoiy  Vigle).  The  absence  of 


S.  aculeatus  from  Santa  Cecilia  (Duell- 
man, 1978)  is  probably  real  rather  than 
sampling  error,  given  the  intensity  of  col- 
lecting over  several  years  at  the  site.  Thus, 
S.  aculeatus  possibly  does  occur  on  the 
right  (south)  bank  of  the  Rio  Napo  and 
will  be  recorded  once  large  collections  are 
made  there. 

The  apparent  geographic  disjunction  of 
Stenocercus  aculeatus  between  northern 
Peru  and  eastern  Ecuador  may  correspond 
to  some  character  differences  among  sam- 
ples that  should  be  studied  more  thor- 
oughly (Cadle,  unpublished  data).  For  ex- 
ample, Peruvian  specimens  of  S.  aculeatus 
have  veiy  deep  postfemoral  pockets  (Type 
5)  in  both  sexes,  whereas  the  postfemoral 
pockets  are  more  weakly  developed  in 
specimens  from  Ecuador  (Type  2  or  3  in 
both  sexes).  Ecuadorian  specimens  also 
appear  to  have  more  scales  in  the  vertebral 
row  and  fewer  subdigital  scales  on  the 
fourth  toe  than  do  Peruvian  specimens.  All 
of  these  impressions  are  based  on  small 
sample  sizes  (Appendix). 

The  significance  of  diese  differences  is 
unclear  without  a  more  detailed  study  of 
variation  among  populations  of  Stenocer- 
cus aculeatus.  However,  one  possibility  is 
that  two  or  more  species  are  represented 
in  specimens  currently  referred  to  S.  acu- 
leatus, in  which  case  the  distributions  of 
individual  taxa  may  be  not  be  contiguous. 
This  is  analogous  to  the  previous  confusion 
of  S.  finihriatus  and  S.  prionotus  with  S. 
aculeatus.  Taxonomic  recognition  of  S. 
finihriatus  and  S.  prionotus  has  concomi- 
tantly reduced  the  geographic  distribution 
understood  for  S.  aculeatus.  Consequently, 
a  more  comprehensive  systematic  analysis 
of  S.  aculeatus  with  special  reference  to  a 
comparison  of  Ecuadorian  and  Peruvian 
populations  is  warranted.  If  two  species 
are  recognized,  the  name  Liocephalus  an- 
gulifer  Werner  (1901)  is  available  for  the^ 
Ecuadorian  populations. 

Key  to  Species  of  the  "Ophryoessoides 
group"  of  Stenocercus 

Because  of  the  general  confusion  about 
the  species  considered  herein  (e.g.,  see  the 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Codle        213 


synonymy  of  Stenocercus  prionohis) ,  I  pro- 
vide the  following  key  as  a  guide  for  iden- 
tifications. The  key  will  work  for  those  spe- 
cies of  Stenocercus  in  Peru  or  Bolivia  with 
keeled  ventral  scales,  enlarged  posterior 
head  scales,  and  one  row  of  moderately  to 
greatly  enlarged  supraoculars  COphryoes- 
soides  group"  as  used  herein).  I  have  also 
included  the  three  other  currently  recog- 
nized species  having  these  characteristics, 
S.  enjthrogaster  (Hallowell),  S.  dumerilii 
(Steindachner),  and  S.  tricristatus  (Du- 
meril),  although  these  are  not  known  from 
Peru  or  Bolivia  and  are  unlikely  to  occur 
there.  Character  and  distributional  data  in 
the  key  for  S.  dunierilii  and  S.  tricristatus 
follow  Avila-Pires  (1995). 

I  also  include  in  the  key  an  undescribed 
species  with  keeled  ventrals  and  enlarged 
head  plates  and  supraoculars  from  Ama- 
zonas  Department,  Peru,  but  I  am  un- 
aware of  other  undescribed  species  of  the 
"Ophrijoessoides  group"  from  Peru  or  Bo- 
livia. However,  an  undescribed  species 
similar  to  Stenocercus  caducus  (but  lacking 
a  posthumeral  flap)  is  known  from  western 
Brazil  (Mato  Grosso)  and  is  not  included 
in  the  key.  Additional  study  of  S.  iridescens 
from  the  Pacfic  lowlands  of  Peru  and  Ec- 
uador is  needed  (Cadle,  1998:  footnote  4) 
and,  as  indicated  above,  a  thorough  mod- 
ern study  of  variation  in  S.  aculeatus  (Am- 
azonian Ecuador  and  Peru)  is  also  war- 
ranted. Other  undescribed  species  may  re- 
side within  either  of  these  named  taxa. 
The  key  Mdll  permit  identification  of  all  Pe- 
ruvian and  Bolivian  taxa  previously  con- 
fused with  S.  aculeatus  (e.g.,  DLxon  and 
Soini,  1975,  1986  [S.  Jimbriatus];  Fugler, 
1983,  1986,  1989  [S.  prionotus])  and  S.  ir- 
idescens (e.g.,  S.  huancahambae  and  S. 
limitaris;  see  Cadle,  1991,  1998).  The  key 
also  should  work  for  Ecuadorian  species, 
with  the  caveat  that  I  have  paid  less  atten- 
tion to  Ecuadorian  Stenocercus  except  as 
necessary  in  conjunction  with  work  on  Pe- 
ruvian species.  Of  the  species  covered, 
only  S.  acideatus,  S.  iridescens,  and  S.  lim- 
itaris are  definitely  known  from  Ecuador. 


The  key  should  be  viewed  as  a  means  of 
identifying  a  set  of  phenotypically  similar, 
but  not  necessarily  closely  related,  species 
within  Stenocercus  in  the  broad  sense.  All 
other  species  of  Stenocercus  in  Peru  and 
Bolivia  have  smooth  (or  at  most  only  very 
weakly  keeled)  ventrals  and  more  frag- 
mented supraoculars  and  head  plates;  see 
Fritts  (1974),  Frost  (1992),  and  Cadle 
(1991,  1998)  for  discussion  and  illustra- 
tions. Many  of  these  species  also  have 
granular  scales  on  the  body  or  posterior 
surface  of  the  thigh,  neither  of  which  is 
present  in  species  covered  by  the  key.  The 
keys  and  discussions  in  Fritts  (1974)  and 
Cadle  (1991,  1998)  are  useful  for  identi- 
fying these  other  species. 

The  key  assumes  familiarity  with  char- 
acters of  the  mite  pockets,  head  scales,  and 
neck  folds  and  crests  outlined  in  Cadle 
(1991)  (see  also  Materials  and  Methods). 
In  most  cases  I  have  used  characters  that 
show  minimal  sexual  dimoqDhism  so  that 
specimens  of  either  sex  can  be  identified; 
exceptions  are  noted.  It  is  useful  to  keep 
in  mind  that,  in  most  species  of  Stenocer- 
cus, scales  of  juveniles  are  more  promi- 
nently keeled  than  in  adults,  even  when 
the  corresponding  scales  of  adults,  such  as 
head  scales  and  dorsal  body  scales,  are 
smooth.  Instances  of  possible  confusion  in 
the  key  are  indicated.  The  extent  of  de- 
velopment of  posthumeral  and  postfemor- 
al  mite  pockets  varies  according  to  sex  and 
size  in  many  species  of  Stenocercus,  al- 
though such  variation  seems  less  extensive 
in  this  set  of  species  than  in  many  others; 
I  have  indicated  the  range  of  variation  in- 
cluding juveniles  and  adults  of  both  sexes 
in  the  key.  Summaiy  geographic  distribu- 
tions are  given  for  each  species  as  a  rough 
guide  to  known  occurrences.  However, 
these  should  be  used  cautiously  as  ancil- 
laiy  information  in  identifying  specimens 
because  distributions  of  species  are  some- 
times poorly  circumscribed.  For  greatest 
utilit}^  the  key  should  be  used  in  conjunc- 
tion with  illustrations  herein  and  in  Cadle 
(1991,  1998)  and  Avila-Pires  (1995). 


214         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


1.  Canthal   and   supraciliaries   forming   a  pro- 

nounced crest  that  ends  in  an  enlarged, 
erect,  postsupraciliaiy  that  may  be  dis- 
tinctly pointed  or  blunt.  Posthumeral  and 
postfemoral  mite  pockets  absent  (Type  1 

in  both  instances)  2 

Cantlials  and  supraciliaries  not  forming  a 
pronounced  crest;  no  enlarged,  erect  post- 
supraciliaiy. Posthumeral  pocket  absent 
(Type  1)  to  deep  (Type  4).  Postfemoral 
pocket  absent  (Type  1)  to  deep  (Type  5) 
3 

2.  Enlarged  postsupraciliary  distinctly  pointed. 

Two  enlarged  scales  above  ear  opening. 
Tibia  approximately  equal  to  thigh  length 

Stenocercus  dumerilii  (Steindachner) 

(northeastern  Para,  Brazil) 
Enlarged  postsupraciliary  blunt.  No  enlarged 
scales  above  ear  opening;  tibia  distinctly 

shorter  than  thigh  

Stenocercus  tricristatus  (Dumeril) 

(known  only  from  the  holotype,  probably 
from  the  state  of  Minas  Gerais,  Brazil) 

3.  Superciliary  scales  projecting  laterally  shelf- 

like above  the  orbit  in  adults,  rectangular 
in  dorsal  view.'^'  Midbody  dorsal  scale  rows 
more  than  55  (59-70).  Postfemoral  pocket 

well  developed  (Type  3  or  5)   

Stenocercus  scapularis  (Boulenger) 

(intermediate  elevations  on  the  Andean 
slopes  of  central  and  southern  Peru;  known 
elevations  greater  than  1,000  m) 
Superciliaiy  scales   not  projecting  laterally, 
the  anterior  ones  elongate,  strongly  over- 
lapping. Midbody  dorsal  scale  rows  fewer 
than  55  (30—53).  Postfemoral  pocket  vari- 
able (Type  1,  2,  3,  or  5)  4 

4.  Posthumeral  mite  pocket  deep  (TjqDe  4)  and 

with  an  associated  scaly  flap  extending 
from  its  anteroventral  border  and  partally 

concealing  it  5 

Posthumeral  mite  pocket  absent  to  deep 
(Type  1,  2,  3,  or  4)  but  without  an  asso- 
ciated flap  6 

5.  Vertebral  crest  strongly  projecting  in  both 

sexes,  serrate,  extending  from  the  nuchal 
region  to  the  proximal  portion  of  the  tail; 
its  individual  scales  triangular,  flaplike. 
Throat  often  with  alternating  oblique  dark 
and  light  stripes.  Two  enlarged,  strongly 
keeled  and  projecting  angulate  temporal 


''  The  superciliaries  in  juveniles  of  Stenocercus  sca- 
pularis have  a  more  typical  shape.  The  extent  of 
shelf-like  projection  and  change  to  a  more  rectan- 
gular shape  seem  positively  correlated  with  body  size 
and  thus  develop  with  age.  The  number  of  dorsal 
scale  rows  and  the  development  of  the  postfemoral 
pocket  are  useful  clues  for  subadults. 


scales  on  each  side  

Stenocercus  prionotiis  Cadle 

(rainforested  lowlands  and  Andean  foothills 
of  eastern  Peru  and  northern  Bolivia) 
Vertebral  crest  low,  nonprojecting  in  both 
sexes  (slightly  higher  in  males),  evident 
mainly  on  neck  and  anterior  body;  its  in- 
dividual scales  prismatic  and  lying  more  or 
less  flat.  Throat  pattern  variable,  but  usu- 
ally consisting  of  light  spots  on  a  darker 
background  when  evident.  Angulate  tem- 
poral scales  keeled,  but  not  greatly  en- 
larged, and  nonprojecting 

Stenocercus  caducus  (Cope) 

(deciduous  woodlands  and  ecotonal  areas  of 

southern  Bolivia,  northern  Argentina,  and 

the  chaco  of  Paraguay) 

6.  A  fringe  of  enlarged  fimbriate  scales  on  the 

distal  posterodorsal  surface  of  thigh.  Sev- 
eral longitudinally  oblique  rows  of  large, 
strongly  keeled  scales  on  shank.'''  Scales 
between  vertebral  and  dorsolateral  crests 
smooth  or  occasionally  veiy  weakly  keeled. 

.-.  Stenocercus  finibriatus  Avila-Pires 

(lowlands  of  eastern  Peru  and  central  western 
Amazonian  Brazil) 
No  fringe  of  fimbriate  scales  on  thigh  or 
strongly  keeled  oblique  scales  on  shank. 
Scales  between  vertebral  and  dorsolateral 
crests  moderately  to  strongly  keeled  at 
least  posteriorly;  dorsolateral  crest  may  be 
weakly  developed,  but  dorsal  scales  still 
strongly  keeled  7 

7.  Posthumeral  mite  pocket  variable  (Type  1,  2, 

3,  or  4).  Postfemoral  mite  pocket  variable 
(Type  1,  2,  3,  or  5).  Head  scales  smooth 
or  keeled.  Angulate  temporal  scales 
keeled,  may  be  projecting  and  bladelike. 
Internasals  usually  4  or  more  (occasionally 
3,  never  2),  often  irregular  in  pattern  and 

shape.  8 

Posthumeral  mite  pocket  absent  or  weakly 
developed  (Type  1  or  2).  Postfemoral  mite 
pocket  absent  (Type  1).  Head  scales 
smooth.  Angulate  temporal  scales  smooth; 
none  bladelike  and  projecting.  Two  polyg- 
onal internasals  in  contact  on  the  midline, 
each  broader  laterally  than  medially. 

Stenocercus  iridescens  (Giinther) 

(Pacific  lowlands  and  intermediate  elevations 
of  western  Ecuador  and  northwestern 

Peru) 

8.  One  to  3  strongly  keeled,  but  nonprojecting, 

angulate  temporal  scales  in  line  with  the 
superciliary  row  between  the  lateral  tem- 


"^  The  fimbriate  scales  form  a  projecting  fringe  on 
the  distal  portion  of  the  thigh.  Both  the  fimbriate 
scales  and  the  oblique  scales  on  the  shank  are  rela- 
tively more  prominent  in  juveniles  than  adults. 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle        215 


porals  and  the  posterior  dorsal  head  scales. 
Two  subequal  can  thai  scales  on  each  side. 

Head  scales  keeled,  at  least  posteriorly 9 

Two  projecting  bladelike  angulate  temporals 
in  line  with  superciliary  row.  A  single  can- 
thai  on  each  side  (rarely,  2  are  present  but 
in  that  case  1  is  much  larger  than  the  oth- 
er). Head  scales  smooth  or  keeled.  11 

9.  Posthumeral  pocket  moderately  developed 

in  males  (Type  2  or  3),  absent  in  females 
(Type  1).  Postfemoral  pocket  absent  in  fe- 
males (Type  1),  moderate  to  deep  in  males 
(Type  3  or  5).  Anterior  gular  scales  weakly 

to  strongly  keeled. 

Stenocercus  erythrogaster  (Hallowell) 

(northern  Colombia) 
Posthumeral  and  postfemoral  pockets  deep 
in  both  sexes  (Types  4  and  5,  respective- 
ly).'' Anterior  gular  scales  smooth  to  weak- 
ly keeled.  10 

10.  Inteiparietal  indistinct,  parietal  eye  not  visi- 

ble. Three  occipitals.  Dark  subocular  bar 
absent.  Three  angulate  temporals  separat- 
ed from  large  posterior  head  scales  by  a 

row  of  tiny  scales Stenocercus  new  species 

(known  from  a  single  specimen  [Appendix] 
from  the  inter-Andean  valley  of  the  Rio 
Maranon  near  Balsas,  Amazonas  Depart- 
ment, Peru) 
Inteiparietal    distinct,    parietal    eye    visible. 
Two  occipitals.   Dark  subocular  bar  pre- 
sent. One  angulate  temporal  much  larger 
than  others  and  in  contact  with  at  least  1 

other  enlarged  posterior  head  scale.  

Stenocercus  liniitaris  Cadle 

(intermediate  elevations  [600—2,200  m]  of  the 
Andes  on  the  Pacific  versant  of  southwest- 
ern Ecuador  and  northwestern  Peru) 

11.  Head  scales  smooth  to  slightly  wrinkled  in 

adults;  weakly  keeled,  wrinkled,  or  rugose 
in  juveniles.  Prominent  dorsolateral  crest 
on  body  from  neck  to  base  of  tail  and  con- 
tinuous with  both  supra-auricular  crest 
and  antehumeral  crest.  Postfemoral  pock- 
et moderate  to  deep  (Type  2,  3,  or  5). 

Stenocercus  aculeatus  (O'Shaughnessy) 

(rainforested  lowlands  and  intermediate  ele- 
vations of  northern  Peru  adjacent  to  the 
Andes  and  in  eastern  Ecuador) 
Head  scales  strongly  keeled  or  multicarinate 
in  juveniles  and  adults.  Dorsolateral  crest, 
when  present,  weak  and  restricted  to  neck 
and    anterior    body.    Postfemoral    pocket 

deep  (Type  5).  

Stenocercus  huancahanibae  Cadle 


''  An  undescribed  species  in  the  ne.xt  couplet  of  the 
key  is  known  only  from  a  single  adult  male.  The  dis- 
tributions of  species  in  couplets  9  and  10  should  be 
used  as  ancillaiy  data  for  identification. 


(diy  inter-Andean  valleys  of  the  upper  Rio 
Maraiion  in  Cajamarca  and  west  central 
Amazonas  departments,  northern  Peru) 

ACKNOWLEDGMENTS 

Loans  and  other  assistance  were  facili- 
tated by  Linda  Ford,  Barrel  Frost,  and 
Charles  W.  Myers  (AMNH);  E.  Nicholas 
Arnold  and  Colin  J.  McCarthy  (BMNH); 
John  Wiens  (CM);  Cassy  Redhed,  Alan 
Resetar,  and  Harold  Voris  (FMNH);  Wil- 
liam E.  Duellman,  Christopher  J.  Raxwor- 
thy,  and  John  E.  Simmons  (KU);  Frank 
Burbrink  and  Douglas  Rossman 
(LSUMNS);  Ross  MacCulloch  and  Robert 
W.  Murphy  (ROM);  Roy  W.  McDiarmid, 
Steven  W.  Gotte,  W.  Ronald  Heyer,  and 
Robert  R  Reynolds  (USNM);  and  Jona- 
than Campbell  and  Michael  B.  Harvey 
(UTA).  Victor  Morales  permitted  me  to  ex- 
amine a  specimen  of  Stenocercus  fimbria- 
tus  in  his  care.  I  am  grateful  to  Wade  C. 
Sherbrooke  for  providing  copies  of  his 
field  notes  and  other  information  on  spec- 
imens he  collected.  I  owe  a  great  debt  to 
the  late  Ernest  E.  Williams,  who  was  ex- 
tremely generous  with  discussion,  notes,  il- 
lustrations, and  encouragement.  Williams, 
Paulo  E.  Vanzolini,  and  Richard  Etheridge 
long  ago  distinguished  Stenocercus  prion- 
otus  and  two  other  species  I  described  (S. 
huancabainbae  and  S.  liniitaris)  but  kindly 
let  my  work  on  the  group  unfold  with  their 
gracious  consent.  Vanzolini  supplied  a 
copy  of  a  portion  of  Balzan  (1931),  helped 
interpret  Balzan's  localities,  and  pointed 
out  Metraux's  work  to  me.  I  am  indebted 
to  several  people  for  the  special  efforts 
they  made  in  tracking  down  information 
about  particular  collections:  Bruce  Patter- 
son (FMNH)  supplied  information  on  the 
collections  of  Colin  Sanborn  and  Hilda 
Heller  from  Puno  Department,  Peru;  Alan 
Resetar  and  Cassy  Redhed  (FMNH)  dug 
into  the  Schmidt  archives  and  found  ad- 
ditional information  about  Hellers  collec- 
tion; Charles  W.  Myers  (AMNH)  did  the 
same  for  Haivey  Basslers  journeys  in 
northern  Peru  and  provided  the  base  map 
used  to  prepare  Figure  1;  Robert  S.  Voss 


216         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


checked  AMNH  sources  for  information 
on  Keays's  Peruvian  localities;  and  Colin  J. 
McCarthy  (BMNH)  clarified  the  confusion 
about  P.  O.  Simons's  "Palca"  locality,  if  not 
the  locality  itself.  Robin  Andrews,  Tom 
Jenssen,  and  A.  Stanley  Rand  discussed  as- 
pects of  geographic  variation  and  the  uses 
of  vertebral  crests  in  S.  prionotus  with  me. 
Laszlo  Meszoly  drew  Figures  4,  5,  and  11. 
For  comments  on  the  manuscript  I  thank 
Richard  Etheridge  and  Charles  W.  Myers. 
The  research  was  supported  in  part  by  a 
faculty  grant  from  the  School  of  Arts  and 
Sciences  of  Harvard  University;  publica- 
tion costs  were  supported  by  the  Colles 
Fund  of  the  MCZ. 

APPENDIX:  SPECIMENS  EXAMINED 

Institutional  abbreviations  are  as  fol- 
lows: 

AMNH  American  Museum  of  Natural 

History,  New  York 

ANSP  Academy  of  Natural  Sciences 

of  Philadelphia 

BMNH  The    Natural    History    Muse- 

um, London 

CM  Carnegie  Museum  of  Natural 

History,  Pittsburgh,  Pennsyl- 
vania 

FMNH  The  Field  Museum,  Chicago 

KU  Natural     History    Museum, 

University  of  Kansas,  Law- 
rence 

LSUMNS  Louisiana  State  University 
Museum  of  Natural  Science, 
Baton  Rouge 

MCZ  Museum  of  Comparative  Zo- 

ology, Harvard  University, 
Cambridge 

ROM  Royal   Ontario   Museum,   To- 

ronto 

USNM  National  Museum  of  Natural 

History,  Washington,  D.C. 

UTA  University  of  Texas  at  Arling- 

ton 

Bracketed  information  was  inferred 
from  sources  listed  in  the  Materials  and 
Methods.  For  specimens  of  other  species 
of  the  "Ophryoessoides  group"  examined 


(S.  huancabamhae,  S.  iridescens,  and  S. 
limitaris),  see  Cadle  (1991,  1998).  Bold- 
face numbered  localities  1—4  are  known  or 
suspected  areas  of  sympatry  between  the 
species  listed  and  Stenocercus  prionotus. 
They  correspond  to  numbered  localities  in 
Figures  1  and  2  and  in  the  text  discussion. 

Stenocercus  aculeatus 

ECUADOR:  Prov.  Morona-Santiago:  Chiguaza 

[ca.  1,000  m;  01°59'S,  77°58'W]  (USNM  200882-84). 
[?Prov.  Napo/Tungurahua]:  Llanganates  area'** 
(FMNH  2.3527).  Prov.  Pastaza:  Rio  Pastaza,  Abita- 
gua  [01°23'S,  78°05'W]  (FMNH  2.5803-05,  26892, 
28011,  28057  [  =  17  specimens]).  Rio  Pastaza,  Alpay- 
acu  [01°28'S,  78°07'W]  (FMNH  3926-27;  MCZ 
8081).  Canelos  [01°35'S,  77°45'W;  530  m]  (MCZ 
38530).  Montalvo,  Rio  Robonaza  [314  m;  02°04'S, 
76°58'W]  (USNM  200892).  [?Prov.  Pastaza]:  Ranos, 
Mera  Trail  [?  =  between  Raiios  and  Mera'^;  approx- 
imately 01°30'S,  78°10'W]  (FMNH  28012). 

PERU:  [Depto.  La  Libertad]:  E  Peru,  Pampa 
Seca,  Rio  MLxiolla  [  =  Rio  Mishollo]  Valley,  Upper 
Huallaga  region,  4300  ft  [2,  1,311  m;  approximately 
08°16'S,  76°58'W]2«  (AMNH  57085).  [Depto.  Lor- 


''  I  have  been  unable  to  localize  this.  The  Cordil- 
lera de  los  Llanganates  is  a  high  range  (to  >4,500  m) 
in  the  Cordillera  Oriental  north  of  the  Rio  Pastaza 
(Paynter,  199.3).  The  locality  may  refer  to  lower  ele- 
vations in  this  range. 

"^  Rafios  is  a  famous  collecting  locality  at  the  foot 
of  Volcan  Tunguraliua  at  1,820  m.  That  would  be  an 
altitudinal  record  for  Stenocercus  aculeatus.  I  inter- 
pret the  locality  as  stated  in  the  FMNH  catalogues 
as  being  on  the  trail  between  Raiios  and  Mera,  which 
is  at  1,160  m.  See  Rrown  (1941)  and  Chapman  (1926) 
for  discussion. 

-"  Hai"vey  Rassler  collected  Stenocercus  for  the 
AMNH  at  two  localities  on  the  Rio  Mixiolla  (  =  Rio 
Mishollo):  Pampa  Seca  and  La  Pinita  (see  Stenocer- 
cus finibriatus),  as  listed  in  AMNH  catalogues.  The 
Rio  Mishollo  originates  in  southeastern  La  Libertad 
Department,  flows  eastward,  and  joins  the  Rio  Hual- 
laga in  southwestern  San  Martin  Department.  The 
elevations  given  for  these  localities,  1,067  m  and 
1,311  m,  indicate  that  they  lie  in  the  narrow  stretch 
of  the  valley  that  straddles  the  boundary  between  La 
Libertad  and  San  Martin  departments  (departmental 
maps  produced  by  the  Instituto  Geografico  Nacional, 
Lima).  I  identify  these  localities  as  Pampaseca  and 
Piiiita,  respectively,  in  extreme  southeastern  La  Lib- 
ertad Department,  as  indexed  by  Stiglich  (1922). 
Roth  localities  are  in  Ongon  District  and  the  coor- 
dinates given  are  those  for  the  town  of  Ongon.  Stig- 
lich (1922)  states  that  Piiiita  is  a  small  village  on  the 
Quebrada  Pedernal,  a  left  tributary  of  the  Rfo  Mish- 
ollo. Apart  from  indicating  that  Pampaseca  is  a  farm. 


New  Species  of  Stenocercus  from  Peru  and  Bolivia  •  Cadle        217 


ETo]:  NE  Peru,  Front  Range  between  Moyabamba 
and  Cahuapanas.  3000  ft  [915  m;  approximately 
05°37'S,  77°00'W]  (AMNH  57083).  Northeastern 
Peru:  Icuta  on  Balsapuerto-Moyabamba  trail,  3500  ft 
[1,067  m;  05°58'S,  76°40'W;  given  as  "Icuto"  or  "len- 
to Cuesta"  by  Lamas,  1976]  (AMNH  56413). 

Stenocercus  caducus 

BOLIVIA:     No    specific    locality:     BMNH 

1946.8.29.76  (holotype  of  Leiocephahis  bolivianus 
Boulenger);  CM  4583-84.  Depto.  Chuquisaca:  Sud 
Cinti,  trail  from  Rinconada  Bufete  to  El  Palmar 
[1,170-2,000  m;  approximately  20°50'S,  64°21'W] 
(UTA  39102).  Depto.  Santa  Cruz:  Buena  Vista,  ca. 
500  m  [17°27'S,  63°40'W]  (MCZ  20625-26,  29023; 
FMNH  16165,  21486,  21511;  BMNH  1927.8.1.163- 
164;  CM  4527,  4550-51,  4558,  4587-88,  4605,  4607, 
4616,  4626,  4634-36,  4641).  Las  Yuntas  [  =  Las  Jun- 
tas;-' 18°38'S,  63°08'W]  (CM  970).  Provincia  Chiqui- 
tos,  Santiago  (Serrania  and  nearby),  700-750  m 
[18°19'S,  59°34'W]  (FMNH  195983).  [Provincial 
Chiquitos,  Canton  El  Cerro,  Finca  Dos  Milanos, 
17°27'30"S,  62°20'00"W  (UTA  38046).  Provincia  Sara, 
eastern  Bolivia,  600  m  [17°27'S,  63°40'W]  (BMNH 
1907.10.31.7-8).  Provincia  Sara,  Santa  Cruz  de  la  Si- 
erra [17°48'S,  63°10'W]  (CM  966,  969,  13018).  Prov- 
incia Sara,  Rio  Surutu  W  of  Buena  Vista  [17°24'S, 
63°51'W]  (CM  4590).  Provincia  Sara,  Rio  Colorado 
[17°38'S,  63°54'W]  (CM  4598).  [Provincia]  Velasco, 
Inselbergs  near  Florida  [14°38'S,  6ri5'W]  (UTA 
38048).  [Depto.  Takija]:  Villa  Monies  [21°15'S, 
63°30'W]  (MCZ  28634).  Misidn  San  Francisco 
[21°15'S,  63°30'W]  (BMNH  98.7.7.5;  specimen  col- 
lected by  Alfredo  Borelli,  whose  San  Francisco  =  Vil- 
la Monies  fide  Paynter,  1992). 

PARAGUAY:  [Depto.  Caaguazu]:  Pastoreo  [ap- 
proximately 25°23'S,  55°52'W]  (MCZ  34214-15). 
[Depto.  Central]:  Asuncion  [25°16'S,  57°40'W] 
(FMNH  9496).  Colonia  Nueva  Italia  [25°37'S, 
57°30'W]  (FMNH  42281). 

Stenocercus  erythrogaster 

COLOMBIA:  [Depto.  Magdalena]:  Rio  Frio 
[30-450  m;  10°55'N,  74°10'W]  (MCZ  29707).  Santa 
Malta  Mountains  [approximately  10°50'N,  73°40'W] 
(MCZ  11303).  Rio  Toribio,  Hacienda  "Papare,"  sec- 
ond river  on  road  from  Cienaga  to  Santa  Marta 
[11°03'N,  74°14'W]  (FMNH  165153).  [Depto.  San- 
tander]:  San  Gil  [1,095  m;  06°33'N,  73°08'W] 
(ANSP  24136,  MCZ  36877). 


Stiglich  (1922)  gives  no  further  information  about  its 
location. 

-'  The  specimen  was  collected  in  November  or  De- 
cember 1913  by  Jose  Steinbach,  who  collected  at  a 
locality  known  as  "Las  Juntas"  during  that  same  pe- 
riod (Paynter,  1992).  The  two  locahties  are  assumed 
to  be  the  same.  The  variant  spelling  "Yuntas"  does 
not  appear  in  any  sources  consulted. 


Stenocercus  fimbriatus 

PERU:  No  specific  locality:  (FMNH  56070). 
Depto.  Huanuco:  ca.  35  km  NE  Tingo  Maria,  Hcda. 
Santa  Elena,  ca.  1000  m  [approximately  08°57'S, 
76°02'W]  (LSUMNS  26966-67).  Approximately  Vi 
mile  E  Universidad  Agraria  de  La  Selva,  Tingo  Maria, 
vicinity  of  Rio  Huallaga  [3,  09°18'S,  75°59'W], 
USNM  193684.  [Depto.  La  Lihertad]:  E  Peru,  La 
Pinita,  Rio  Mixiolla,  tributaiy  of  upper  [Rio]  Hualla- 
ga, 3500  ft  [1,067  m;  approximately  08°16'S,  76°58'W; 
see  footnote  20]  (AMNH  56797-98).  [Depto.  Lor- 
ETO]:  E  Peru,  Contamana,  Ucayali  River  valley  [134 
m;  07°15'S,  74°54'W]  (AMNH  56803).  E  Peru,  E  of 
Contamana  on  trail  to  Contaya,  700  ft  [213  m;  ap- 
proximately 07°15'S,  74°54'W]  (AMNH  56781-82).  E 
Peru,  Pampa  Hermosa,  mouth  of  Rio  Cushabatay, 
500  ft  [152  m]  [1,  07°12'S,  75°17'W]  (AMNH  56788, 
56790-92,  56794-96,  56801-02).  Mishana,  Rio  Na- 
nay,  Estacion  Biologica  Cauicebus,  150  m  [03°53'S, 
73°27'W]  (USNM  "222377).  Mishuana  [  =  Mishana; 
150  m,  03°53'S,  73°27'W]  (KU  212628).  Depto.  Ma- 
DREDE  Dios:  Pakitza  Station  [Rio  Manu],  Manu  Na- 
tional Park  [4,  11°56'S,  71°17'W]  (Victor  R.  Morales 
18235).  Depto.  Ucayali:  Rio  Curanja,  Balta,  ap- 
proximately 300  m  [approximately  10°08'S,  71°13'W] 
(LSUMNS  17519,  25402-04,  26720-23).  Alto  [Rio] 
Purris,  Alto  [Rfo]  Curanja,  Igarape  Champuiaco 
[9°34'S,  70°36'W]  (MCZ  61226)"  Peru/Brazil  frontier, 
Utoquinia  Region,  1000  ft.  [305  m;  approximately 
08°00'S,  74°00'W]22  (AMNH  56789,  56799-800). 

Stenocercus  scapulari^^ 

PERU:  No  specific  locality:  (FMNH  56444). 
[Depto.  Junin]:  Chanchamayo,  1200  m  [approxi- 
mately 11°03'S,  75°47'W]  (FMNH  40608-11).  Pere- 
ne,  1200  m  [10°58'S,  75°13'W]  (MCZ  49580-81). 
Tarma,  Chanchamayo,  1300  m  [11°25'S,  75°42'W] 
(FMNH  45522).  [Depto.  Puno]:  Sagrario,  Rio  Qui- 
tun    [approximately    1,020    m;    13°55'S,    69°41'W] 


--  The  region  referred  to  is  north  to  northeast  of 
Pucallpa.  The  variant  spellings  Utoquinia,  Utoquinea, 
and  Uroquinea  are  in  the  literature  and  are  applied 
to  a  right-bank  tributary  of  the  Rio  Ucayali,  a  village 
on  the  Rio  Ucayali,  and  an  airstrip  on  the  Rio  Uto- 
quinia near  the  Brazilian  border.  The  entire  region  is 
less  than  500  m  in  elevation  except  for  a  small  raised 
area  near  the  Brazilian  border  that  attains  nearly  800 
m  and  that  is  apparently  the  source  of  the  Rio  Uto- 
quinia. 

^^  The  occurrence  of  Stenocercus  scapularis  at  Rur- 
renabaque.  El  Beni  Department,  Bolivia,  as  reported 
for  two  specimens  in  the  AMNH  (Burt  and  Burt, 
1931:  273)  is  apparently  based  on  a  misidentification. 
These  specimens  are  probably  either  S.  prionotus 
(most  likely)  or  S.  caducus  (see  Distribution  Patterns 
in  Stenocercus  prionotus). 


218         Bulletin  Museum  of  Comparative  Zoology,  Vol.  157,  No.  3 


(FMNH  40408).  "Camp  4"  [between  Santo  Domingo 
and  La  Pampa;  approximately  13°44'S,  69°37'W]-^ 
(FMNH  40409).  Juliaca,  Lake  Aracona,  16,600  ft. 
[shipping  point  only;  correct  locality  is  on  the  right 
laank  of  the  Rio  Inambari,  1,830  m,  13°30'S, 
70°00'W]^'^  (AMNH  1701). 


-^  According  to  the  field  catalogue  in  the  FMNH 
Mammal  Di\dsion  the  collector,  Colin  Sanborn,  was 
in  Santo  Domingo  on  20  October  1941  and  in  La 
Pampa  on  23  October  (see  also  notes  in  Stephens  and 
Traylor,  1983).  The  specimen  FMNH  40409  was  col- 
lected 21  October,  and  thus  "Camp  4"  is  assumed  to 
be  between  these  points. 

-''The  specimen  was  collected  in  1900  by  H.  H. 
Keays,  who  collected  many  mammals  and  other  ver- 
tebrates in  southern  Peru,  primarily  for  the  American 
Museum  of  Natural  History.  It  is  clear  that  most  of 
the  specimens  labeled  with  the  locality  "Juliaca"  (a 
town  on  the  Peruvian  altiplano  near  Lake  Titicaca) 
actually  came  from  farther  north  in  the  Rio  Inambari 
valley.  Allen  (1900:  219;  1901:  41)  provides  the  fol- 
lowing information: 

The  Museum  has  recently  received  two  small  col- 
lections of  mammals  made  by  Mr.  H.  H.  Keays,  at 
Juliaca,  in  southeatern  Peru,  a  little  to  the  west- 
ward of  Lake  Titicaca.  Mr  Keays  writes:  "Our 
camp  is  situated  in  the  loop  of  the  Inambaiy  River. 
The  countiy  is  very  broken,  with  deep  narrow  can- 
ons, and  is  covered  with  a  dense  undergrowth  oi 
shrubs  and  vines,  with  here  or  there  a  palmetto  or 
a  cedar  rising  above  the  surrounding  vegetation." 
He  gives  the" altitude  as  6000  feet  [1,830  m],  and 
the  position  as  latitude  13°30'  S.,  longitude  70°  W. 

...  it  is  necessary  to  correct  a  misleading  statement 
in  my  former  paper  in  respect  to  the  locality  where 
the  .  .  .  collections  were  made.  Mr.  Keays's  post- 
office  address  was  Juliaca,  and  through  lack  of  ex- 
plicit information,  it  was  inferred  that  the  Inca 
Mines,  where  he  collected,  were  in  the  immediate 
vicinity  of  Juhaca  .  .  .  the  Inca  Mines  are  situated 
about  200  miles  northeast  of  JuHaca,  on  the  east 
side  of  the  Andes,  on  the  Inambaiy  River,  a  trib- 
utaiy  of  the  Amazon,  and  at  a  much  lower  altitude 
than  Juliaca.  The  altitude  and  geographical  position 
were  correctly  given  in  the  former  paper,  but  in 
place  of  Juliaca,  .  .  .  read  Inca  Mines. 

Keays's  information  quoted  by  Allen  places  the  local- 
ity on  the  right  bank  of  the  Rio  Inambari  in  the  foot- 
hills of  an  outlying  Andean  spur  separating  the  Rio 
Inambari  from  upper  tributaries  of  the  Rio  Tambo- 
pata.  I  have  not  located  a  Lake  Aracona  and  suspect 
that  this  is  an  error  for  Lake  Aricoma,  a  high  Andean 
lake  on  tlie  route  between  Juliaca  and  the  location  of 
Keays's  camp.  However,  it  is  not  at  all  clear  why  this 
name  is  associated  with  the  locality.  No  notes  or  cor- 
respondence of  Keays  are  in  the  AMNH  mammal 
department  archives  for  further  clarification  (R.   S. 


Stenocercus  sp. 

PERU:   Depto.  Amazonas:   17  km  ENE   Balsas 
[06°49'S,  7S°00'W]  (ROM  16458). 

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